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Purpose This paper aims to examine the role of individual action in addressing challenges of sustainability, and to help marketing scholars and practitioners better understand what motivates sustainable living. Design/methodology/approach Semistructured interviews with 35 individuals self-identifying as sustainable shed light on motivations and identity expression in sustainable living. Four Drive Theory, and Personal and Social Identity Theory (operationalized through the Dynamic Model of Identity Development), provide this study’s guiding theoretical framework. Data analysis was informed by the Gioia methodology. Findings Individuals differently express their personal and social identities through sustainable living, and are differently motivated to live sustainably. Those expressing personal identity salience through sustainable living draw on a broader set of motivations than those expressing social identity salience. This results in varying levels of commitment to sustainable living, with differences also found in individuals’ personal satisfaction derived from their sustainable living efforts. Based on these findings, a novel typology of sustainable individuals is developed. Research limitations/implications This study is limited by its focus on one geographic area and relatively small sample size. A key implication is the need to consider both personal and social identity when studying behavior in other marketing contexts. Practical implications The research provides important insights for marketing practitioners, policymakers and others seeking to better categorize sustainable individuals and target marketing messages to encourage sustainable behaviors. Originality/value This paper contributes to marketing scholarship by providing new insights on the role of identity and motivations in sustainable living. It introduces a novel typology of sustainable individuals, founded on differences in identity expression and motivational drives, which are also associated with the range of sustainable behaviors people engage with and how individuals make sense of these behaviors.

International Journal of Biological Macromolecul

Gyrocotyle discoveryi n. sp. Type-host: Hydrolagus mirabilis (Collett) (Chimaeriformes: Chimaeridae), large-eyed rabbitfish. Type-locality: Goban Spur, North-East Atlantic. Other localities: Porcupine Seabight (51��09 0 N, 11��55 0 W, depth 1,200 m, 30.xi.2002, RRS Discovery Cruise, No. 15048-14, 15); Goban Spur (49��49 0 N, 11��44 0 W, depth 1,175���1,250 m, 27.iv.2001, RRS Discovery Cruise 252, No. 13963/17, 20, 24, 72; 49��41 0 N, 11��53 0 W, depth 1,053���1,077 m, 23.iv.2001, RRS Discovery Cruise 252, No. 13962/4; 49��47 0 N, 11��58 0 W, depth 1,240���1,360 m, 19.x.2002, RRS Discovery Cruise D 266, No. 15066-124, 125; 51��09 0 N, 11��55 0 W, depth 1200 m, 30.ix.2002, RRS Discovery Cruise D 266, No. 15063-103), North-East Atlantic. Type-material: Holotype (NHMUK 2019.11.21.3), paratypes (NHMUK.2019.11.21.4-13 from Goban Spur; NHMUK.2019.11.21.14-19 from Porcupine Sea Bight).). Site in host: Spiral intestine. Representative DNA sequences: MN655879 and MN655881 (ssrDNA); MN657003-MN657005, MN657007- MN657009, MN657011 (lsrDNA, domains D1-D3). ZooBank registration: The Life Science Identifier (LSID) for Gyrocotyle discoveryi n. sp. is urn:lsid:- zoobank.org:act:7B028A0B-B8EB-495E-A9F1- F29DDB60B89A. Etymology: The species is named after the RRS Discovery, the NERC research vessel on which the specimens were collected. Description [Based on 17 specimens; Figs. 4���6.] With characters of the order. Body relatively squat, with deeply crenulated margins, 8,634 ���17,586 X 4,996 ���8,439 (12,071 X 6,676), width 36���98 (60)% of length. Anterior sucker distinct, 1,149 ���1,621 (1,347) long, 596���1,022 (771) wide. Uterus large, in central part of body, 1,546 ���2,996 (2,232) from anterior extremity, 2,453 ���6,226 (4,533) long, 35���43 (38)% of body length. Rosette distinct, fairly complex, 1,797 ���3,092 (2,575) long, junction with soma not clear. Eggs tanned, operculate, 85��� 97 X 39���56 (89 X 49). Diagnosis Gyrocotyle discoveryi n. sp. can be diagnosed from other congeners on the basis of unique nucleotide characters in our rDNA alignments (listed as alignment position-nucleotide): ssrDNA: 176-G, 782-G, 862-G, 973-C; lsrDNA: 573-A, 800-T, 1,245-C, 1,246- C, 1,247-G, 1,360-T, 1,369-T, 1,375-C, 1,379-G, 1,382-G, 1,391-A, 1,449-T, 1,468-T, 1,477-T. Remarks Mauchline & Gordon (1984) reported a ������cestode������ in Hy. mirabilis from the Rockall Trough off NW Scotland, which is, as far as we are aware, the only possible record of a Gyrocotyle from this host. Two species of Gyrocotyle, G. major van der Land & Templeman, 1968 and G. abyssicola van der Land & Templeman, 1968, have been reported from its congener, the small-eyed rabbit fish Hydrolagus affinis (de Brito Capello) on the edges of the continental shelf off the eastern coast of Newfoundland (van der Land & Templeman, 1968). These two species are illustrated as much more elongate than our specimens, with less complex lateral wrinkling and small rosettes. The worms were recovered from frozen hosts, so the gross morphology may well not be of significance in differentiating these species. Subsequently, these two taxa have been reported from the same host species off south-western Greenland by Karlsbakk et al. (2002) and, puzzlingly, from a rhinochimaerine species, the straight-nosed rabbit fish Rhinochimaera atlantica Holt & Byrne, off the Scotian Shelf by Hogans & Hurlbut (1984). In the North-East Atlantic, R. atlantica has not been found harbouring Gyrocotyle, but it does harbour the strobilate tapeworm Chimaerocestos prudhoei Williams & Bray, 1984 and a congeneric host, the Pacific spookfish R. pacifica (Mitsukuri, 1895) also harbours a species of Chimaerocestos Williams & Bray, 1984 (see Caira et al., 1999, 2014). Other records of Gyrocotyle spp. from Hydrolagus spp. are from the Pacific Ocean (see Bandoni & Brooks, 1987). There are no reliable morphological characters to differentiate this species or indeed any of the gyrocotylidean species circumscribed by molecular means. Therefore, the species is diagnosed by its relatively marked sequence divergence from those of recognised species., Published as part of Bray, Rodney A., Waeschenbach, Andrea, Littlewood, D. Timothy J., Halvorsen, Odd & Olson, Peter D., 2020, Molecular circumscription of new species of Gyrocotyle Diesing, 1850 (Cestoda) from deep-sea chimaeriform holocephalans in the North Atlantic, pp. 285-296 in Systematic Parasitology 97 on pages 292-293, DOI: 10.1007/s11230-020-09912-w, http://zenodo.org/record/4623814, {"references":["Mauchline, J., & Gordon, J. D. M. (1984). Incidence of parasitic worms in stomachs of pelagic and demersal fish of the Rockall Trough, northeastern Atlantic Ocean. Journal of Fish Biology, 24, 281 - 285.","van der Land, J., & Templeman, W. (1968). Two new species of Gyrocotyle (Monogenea) from Hydrolagus affinis (Brito Capello) (Holocephali). Journal of the Fisheries Research Board of Canada, 11, 2365 - 2385.","Karlsbakk, E., Aspholm, P. E., Berg, V., Hareide, N. R., & Berland, B. (2002). Some parasites of the small-eyed rabbitfish, Hydrolagus affinis (Capello, 1867) (Holocephali), caught in deep waters off SW Greenland. Sarsia, 87, 179 - 184.","Hogans, W. E., & Hurlbut, T. R. (1984). Parasites of the knifenose chimaera, Rhinochimaera atlantica, from the northwest Atlantic Ocean. Canadian Field-Naturalist, 98, 365.","Williams, H. H., & Bray, R. A. (1984). Chimaerocestos prudhoei gen. et sp. nov., representing a new family of tetraphyllideans and the first record of strobilate tapeworms from a holocephalan. Parasitology, 88, 105 - 116.","Caira, J. N., Jensen, K., & Healy, C. J. (1999). On the phylogenetic relationships among tetraphyllidean, lecanicephalidean and diphyllidean tapeworm genera. Systematic Parasitology, 42, 77 - 151.","Caira, J. N., Jensen, K., Waeschenbach, A., Olson, P. D., & Littlewood, D. T. J. (2014). Orders out of chaos - molecular phylogenetics reveals the complexity of shark and stingray tapeworm relationships. International Journal for Parasitology, 44, 55 - 73.","Bandoni, S. M., & Brooks, D. R. (1987). Revision and phylogenetic analysis of the Gyrocotylidea Poche, 1926 (Platyhelminthes: Cercomeria: Cercomeromorpha). Canadian Journal of Zoology, 65, 2369 - 2389."]}

Gyrocotyle haffii n. sp. Type-host: Harriotta raleighana Goode & Bean (Chimaeriformes: Rhinochimaeridae), bent-nosed chimaera. Type-locality: Goban Spur (49°46 0 N, 12°21 0 W, depth 1,631–1,653 m, 22-23.iv.2001; RRS Discovery Cruise 252, No. 13951/14), North-East Atlantic. Type-material: Holotype (NHMUK. 2019.11.21.1), paratype (NHMUK. 2019.11.21.2). Site in host: Spiral intestine. Representative DNA sequences: MN655880 (ssrDNA); MN657006 (lsrDNA, domains D1-D3). ZooBank registration: The Life Science Identifier (LSID) for Gyrocotyle haffii n. sp. is urn:lsid:- zoobank.org:act:7717F9D6-4C9D-4C59-8D0A- 4C5E306B4671. Etymology: The species is named in honour of our late colleague and friend Professor Harford ‘Haffi 0 Williams in recognition of his contribution to the understanding of the Gyrocotylidea. Description [Based on a single intact, immature whole worm and second immature worm from which the central portion had been excised for molecular analysis; Figs. 2, 3]. With characters of the order. Body elongate with minute annular ridges; no large lateral flap. Length 23,504; greatest width near anterior extremity, 3,131. Rosette relatively small with few crenulations, 1,943 long. Anterior sucker large, oval, 1,750 X 1,223. Reproductive system immature; anlagen commences 2,852 from anterior extremity, 8,191 long; consisting of a long, narrow patch of stained tissue reaching, and a branched section passing, towards lateral margin of worm; apparently opening at c. 268 from anterior extremity. Only other evidence of reproductive organs is putative vitelline glands scattered around posterior extremity of anlagen. Diagnosis Gyrocotyle haffii n. sp. can be diagnosed from other congeners on the basis of unique nucleotide characters in our rDNA alignments (listed as alignment positionnucleotide): ssrDNA: 218-T, 723-A, 746-C, 747-A, 748-G, 1,158-T, 1,654-G, 1,673-C, 2,115-C; lsrDNA: 612-T, 837-A, 875-T, 1,306-A, 1,395-C, 1,402-A, 1,501-G. Remarks As far as we are aware there is only one previous report of a gyrocotylidean from Ha. raleighana, the bentnose chimaera. Parukhin (1966) reported ‘‘ Gyrocotyloides nybelini Fuhrmann, 1931 ’’ in this host from the South Atlantic Ocean. Parukhin (1968) repeated this report saying (in translation) ‘‘Found in Callorhynchus capensis. Two adult parasites were found in two fish. In addition, six larvae were found in one of them. In addition to C. capensis, specimens were found in two Hariota [sic] raleighana. In both cases there were two specimens. Previously, this species was observed in the Atlantic in Chimaera monstrosa ’’. In addition, it seems likely that the records of ‘cestode adults 0 from Ha. raleighana, Hy. mirabilis and C. monstrosa from the Rockall Trough off NW Scotland by Mauchline & Gordon (1984) refer to Gyrocotyle spp. There is no reliable morphological character to differentiate this species or indeed any of the gyrocotylidean species circumscribed by molecular means. Therefore, the species is diagnosed by its relatively marked sequence divergence from those of recognised species.

Gyrocotyle discoveryi n. sp. Type-host: Hydrolagus mirabilis (Collett) (Chimaeriformes: Chimaeridae), large-eyed rabbitfish. Type-locality: Goban Spur, North-East Atlantic. Other localities: Porcupine Seabight (51°09 0 N, 11°55 0 W, depth 1,200 m, 30.xi.2002, RRS Discovery Cruise, No. 15048-14, 15); Goban Spur (49°49 0 N, 11°44 0 W, depth 1,175–1,250 m, 27.iv.2001, RRS Discovery Cruise 252, No. 13963/17, 20, 24, 72; 49°41 0 N, 11°53 0 W, depth 1,053–1,077 m, 23.iv.2001, RRS Discovery Cruise 252, No. 13962/4; 49°47 0 N, 11°58 0 W, depth 1,240–1,360 m, 19.x.2002, RRS Discovery Cruise D 266, No. 15066-124, 125; 51°09 0 N, 11°55 0 W, depth 1200 m, 30.ix.2002, RRS Discovery Cruise D 266, No. 15063-103), North-East Atlantic. Type-material: Holotype (NHMUK 2019.11.21.3), paratypes (NHMUK.2019.11.21.4-13 from Goban Spur; NHMUK.2019.11.21.14-19 from Porcupine Sea Bight).). Site in host: Spiral intestine. Representative DNA sequences: MN655879 and MN655881 (ssrDNA); MN657003-MN657005, MN657007- MN657009, MN657011 (lsrDNA, domains D1-D3). ZooBank registration: The Life Science Identifier (LSID) for Gyrocotyle discoveryi n. sp. is urn:lsid:- zoobank.org:act:7B028A0B-B8EB-495E-A9F1- F29DDB60B89A. Etymology: The species is named after the RRS Discovery, the NERC research vessel on which the specimens were collected. Description [Based on 17 specimens; Figs. 4–6.] With characters of the order. Body relatively squat, with deeply crenulated margins, 8,634 –17,586 X 4,996 –8,439 (12,071 X 6,676), width 36–98 (60)% of length. Anterior sucker distinct, 1,149 –1,621 (1,347) long, 596–1,022 (771) wide. Uterus large, in central part of body, 1,546 –2,996 (2,232) from anterior extremity, 2,453 –6,226 (4,533) long, 35–43 (38)% of body length. Rosette distinct, fairly complex, 1,797 –3,092 (2,575) long, junction with soma not clear. Eggs tanned, operculate, 85– 97 X 39–56 (89 X 49). Diagnosis Gyrocotyle discoveryi n. sp. can be diagnosed from other congeners on the basis of unique nucleotide characters in our rDNA alignments (listed as alignment position-nucleotide): ssrDNA: 176-G, 782-G, 862-G, 973-C; lsrDNA: 573-A, 800-T, 1,245-C, 1,246- C, 1,247-G, 1,360-T, 1,369-T, 1,375-C, 1,379-G, 1,382-G, 1,391-A, 1,449-T, 1,468-T, 1,477-T. Remarks Mauchline & Gordon (1984) reported a ‘‘cestode’’ in Hy. mirabilis from the Rockall Trough off NW Scotland, which is, as far as we are aware, the only possible record of a Gyrocotyle from this host. Two species of Gyrocotyle, G. major van der Land & Templeman, 1968 and G. abyssicola van der Land & Templeman, 1968, have been reported from its congener, the small-eyed rabbit fish Hydrolagus affinis (de Brito Capello) on the edges of the continental shelf off the eastern coast of Newfoundland (van der Land & Templeman, 1968). These two species are illustrated as much more elongate than our specimens, with less complex lateral wrinkling and small rosettes. The worms were recovered from frozen hosts, so the gross morphology may well not be of significance in differentiating these species. Subsequently, these two taxa have been reported from the same host species off south-western Greenland by Karlsbakk et al. (2002) and, puzzlingly, from a rhinochimaerine species, the straight-nosed rabbit fish Rhinochimaera atlantica Holt & Byrne, off the Scotian Shelf by Hogans & Hurlbut (1984). In the North-East Atlantic, R. atlantica has not been found harbouring Gyrocotyle, but it does harbour the strobilate tapeworm Chimaerocestos prudhoei Williams & Bray, 1984 and a congeneric host, the Pacific spookfish R. pacifica (Mitsukuri, 1895) also harbours a species of Chimaerocestos Williams & Bray, 1984 (see Caira et al., 1999, 2014). Other records of Gyrocotyle spp. from Hydrolagus spp. are from the Pacific Ocean (see Bandoni & Brooks, 1987). There are no reliable morphological characters to differentiate this species or indeed any of the gyrocotylidean species circumscribed by molecular means. Therefore, the species is diagnosed by its relatively marked sequence divergence from those of recognised species.

Genus Biospeedotrema n. g. urn:lsid:zoobank.org:act: 6 A 819058 -BDC 2-4752 -A 4 DB- 9 A 47 A 5 EC 8 BDD Diagnosis. Body small, broadly pyriform to distinctly wider than long. Tegument unarmed, but may have annular wrinkles. Oral sucker subglobular to transversely oval, subterminal to terminal. Ventral sucker rounded to oval, larger than oral sucker, just pre-equatorial. Prepharynx short, dorsal to oral sucker. Pharynx large, subglobular to oval. Oesophagus distinct. Intestinal bifurcation about equatorial, dorsal to ventral sucker. Caeca blind, wide, divergent, reaching almost to or just past testes, not to posterior extremity. Testes two, oval, pyriform, may be lobate, symmetrical to slightly oblique, in anterior hindbody. Cirrus sac claviform, reaches dorsally to ventral sucker. Internal seminal vesicle saccular. Pars prostatica long, narrow, surrounded by gland-cells or indistinct. Ejaculatory duct short to long. Genital atrium distinct. Genital pore median to slightly dextrally submedian, immediately anterior to ventral sucker. Ovary subglobular, oval or subtriangular, just anterior to or overlapping right testis, overlapping ventral sucker. Seminal receptacle uterine. Laurer’s canal opens dorsally at about level of posterior margin in ventral sucker or anterior hindbody. Mehlis’ gland ventral to ovary. Uterus reaching between testes, or distinctly into post-testicular region. Eggs large, tanned, operculate, relatively few. Metraterm not distinct. Vitellarium follicular, in lateral fields from pharynx or oral sucker to just into post-testicular region, mainly lateral to caeca. Excretory pore terminal. Vesicle short, flattens into broad vesicle just posterior to testes. Type species Biospeedotrema jolliveti n. sp. Etymology. This new genus is named after the Biospeedo expedition on which the material was collected. Remarks. Biospeedotrema n. gen. is placed in the subfamily Stenakrinae as its members have a welldeveloped muscular cirrus sac and a uterine seminal receptacle. Cribb (2005) recognised six genera in the subfamily. The ovary is usually more or less globular to oval, but can be subtriangular. Utilising the key in Cribb (2005) the closest genus is Caudotestis Issaitschikov, 1928 (see below), but Biospeedotrema differs in the more or less symmetrical testicular configuration, with the uterus passing between the testes, sometimes distinctly into the post-testicular region. Of the four species of Stenakron Stafford, 1904 recognised by Cribb (2005), only one has symmetrical testes, namely Stenakron vitellosum (Manter, 1934) from the shortbeard codling Laemonema barbatulum Goode & Bean (Gadiformes, Moridae) in 538 +m depth (294 + fathoms) off Tortugas, Florida (Manter 1934). It has a trilobed ovary (a characteristic of the genus) and the uterus does not extend posteriorly to the testes. Biospeedotrema, Caudotestis and Stenakron are mainly parasites of deep-sea fishes (Klimpel et al. 2009).

Biospeedotrema jolliveti n. sp. (Fig. 1) urn:lsid:zoobank.org:act: 5 F 4 B 8936 -D 162 - 4 F 77 - 9 D 80-240 E 32 DCAD 02 Type host. Ventichthys biospeedoi Neilsen, M��ller & Segonzac (Ophidioformes: Ophidiidae), East-Pacific ventbrotula. Site. ? intestine (not given) Type locality. South East Pacific Rise, hydrothermal vent site Oasis, 17 �� 25.38 ���S, 113 �� 12.29 ���W; Biospeedo cruise, submersible Nautile, R/V Atalante, dive PL 1582, baited trap B 2, depth 2586 m, 22 April 2004, Biospeedo labels: 1582 ���2, 1582��� 5. Type specimens. MNHN holotype 1 slide MNHN HEL 373; paratypes 3 wholemount slides MNHN HEL 374 375 376, 5 section slides MNHN HEL 377 378 379 380 381; BMNH paratypes 2013.12.3.4- 6. GenBank numbers. rDNA KF 733985, rDNA KF 733988. Etymology. This species named after Dr. Didier Jollivet who led the Biospeedo expedition. Description. Based on 7 whole-mounts and 2 sets of serial sections specimens. Measurements and ratios in Table 1. Body small, broadly pyriform (Fig. 1). Tegument unarmed. Oral sucker subglobular, subterminal. Ventral sucker rounded to oval, much larger than oral sucker, centre just pre-equatorial. Prepharynx short, dorsal to oral sucker. Pharynx large, subglobular. Oesophagus distinct, thin-walled, long, sigmoid. Intestinal bifurcation about equatorial, dorsal to ventral sucker. Caeca blind, wide, divergent, reaching just past testes, not to posterior extremity. Testes two, oval to pyriform, transversely elongate, symmetrical to slightly oblique, in anterior hindbody. Cirrus sac claviform, reaches to about mid-ventral sucker. Internal seminal vesicle saccular. Pars prostatica long, narrow, surrounded by gland-cells. Ejaculatory duct short. Genital atrium distinct. Genital pore median to slightly dextrally submedian, immediately anterior to ventral sucker. Ovary oval to subtriangular, just anterior to or overlapping right testis, overlapping ventral sucker. Details of female proximal system disrupted in sections by eggs. Seminal receptacle uterine. Laurer���s canal opens dorsally at about level of posterior margin in ventral sucker. Mehlis��� gland ventral to ovary. Uterus reaching between testes, or just into post-testicular region, then passes sinistrally over dorsal surface of ventral sucker. Eggs large, tanned, operculate, relatively few. Metraterm not distinct. Vitellarium follicular, in lateral fields from pharynx or oral sucker to just into post-testicular region, narrow band confluent or approaching in region of testes, mainly lateral to caeca, but encroaching dorsally. Excretory pore terminal, in slight notch. Vesicle short, flattens into broad vesicle just posterior to testes. Remarks. The features differentiating this species from its congeners are discussed below., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on pages 75-76, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628

Biospeedotrema biospeedoi n. sp. (Fig. 3) urn:lsid:zoobank.org:act: 8381 CFB 8 - 5 EA 9 - 4 FD 7 -A 9 DE-A 161 B 7 C0E 664 Type host. Thermichthys hollisi (Cohen, Rosenblatt & Moser) (Ophididiiformes: Bythitidae ), Hollis brosmekvabbe. Site. Small and large intestine. Type locality. South East Pacific Rise, hydrothermal vent site Hobbs, 17 ° 35.20 ’S, 113 ° 14.75 ’W; Biospeedo cruise, PL 1588, depth 2598 m, baited trap, 29 Apr. Biospeedo labels: 1588 –3, 1588–6, 1588– 8. Type specimens. MNHN holotype 1 slide MNHN HEL 391, paratypes 6 wholemount slides MNHN HEL 392 393 394 395 396 397, 4 section slides. MNHN HEL 398 399 400 401; BMNH paratypes 2013.12.3.7- 10. GenBank number. rDNA KF 733986. TABLE 1. Measurements and ratios of digeneans of hydrothermal vent fishes. Width as % of body-length 39.3–51.1 (43.6) 57.7–82.2 (67.9) 60.6 104–133 (121) 47.6–85.6 (71.6) 22.5–53.9 (37.2) …… continued on the next page p TABLE 1. (Continued) Ventral sucker to ovary as % of body- 2.20–17.8 (7.41) 0–2.11 (0.40) 0 0–7.68 (2.21) 0–6.85 (1.88) 0–2.27 (0.75) length Etymology. This species is named after the Biospeedo expedition on which the material was collected Description. Based on 11 wholemounts and 2 sets of serial sections. Measurements and ratios in Table 1. Body small, oval to fusiform, widest at level of ventral sucker (Fig. 3). Tegument unarmed, but thrown into numerous denticulate annular folds. Oral sucker shallow cup-shaped, almost terminal. Ventral sucker rounded, larger than oral sucker, just pre-equatorial. Prepharynx short. Pharynx oval. Oesophagus long, thin-walled. Intestinal bifurcation dorsal to ventral sucker. Caeca blind, divergent, narrow, reaching just into hindbody or just to testis. Testes lobate, symmetrical, separated by uterus, in anterior hindbody. Cirrus sac elongate claviform, reaching into hindbody, proximal extremity in sinistral anterior hindbody, curves dorsally to ventral sucker. Internal seminal vesicle saccular. Pars prostatica narrow, surrounded by gland-cells. Ejaculatory duct? Genital atrium small. Genital pore median to slightly submedian, just anterior to ventral sucker. Ovary oval, between dextral testis and cirrus sac. Seminal receptacle uterine. Laurer’s canal opens dorsally in mid post-testicular region. Mehlis’ gland not clearly seen. Uterus mainly post-testicular but also intertesticular. Eggs large, operculate, tanned, few. Metraterm not clearly differentiated. Vitellarium follicular, follicles small, irregular, lateral fields reaching from anterior part of ventral sucker to anterior part of testes, confluent in narrow band dorsally to ventral sucker, lateral and dorsal to caeca. Excretory pore terminal, to dorsally subterminal. Vesicle I-shaped, reaches between testes to ovary. Remarks. In basic morphology this species is similar to B. jolliveti, but differs in being generally smaller. The vitelline fields are shorter, filling a smaller proportion of the body-length, and the cirrus sac is relatively larger (Table 1). The uterus tends to extend further posteriorly, relative to the testes. The morphological similarity between B. jolliveti and this species is also reflected in the molecular data; 1300 base pairs of lsrDNA differ only in two positions. Biospeedotrema parajolliveti differs in body-shape and the ratios of vitelline field length and ventral sucker size to body-length. The uterus tends to extend further posteriorly, relative to the testes.

Biospeedotrema parajolliveti n. sp. (Figs. 2) urn:lsid:zoobank.org:act: 4 B 22 B 98 B- 50 F 4 - 4 D 19 -AA 2 D- 7 D 47776 CCD 29 Type host. Thermichthys hollisi (Cohen, Rosenblatt & Moser) (Ophididiiformes: Bythitidae ), Hollis brosmekvabbe. Site. Large intestine and gall-bladder. Type locality. South East Pacific Rise, hydrothermal vent site Hobbs, 17 �� 35.20 ���S, 113 �� 14.75 ���W; Biospeedo cruise, PL 1588, depth 2598 m, baited trap, 29 Apr. Biospeedo labels: 1588 ���1, 1588��� 3. Type specimens. MNHN holotype 1 slide MNHN HEL 382, paratypes 4 wholemount slides, MNHN HEL 383 384 385 386, 4 section slides MNHN HEL 387 388 389 390; BMNH paratypes 2013.12.3.1- 3. Etymology. This species is named to emphasise its similarity to Biospeedotrema jolliveti. Description. Based on 8 wholemount specimens and 2 sets of serial sections. Measurements and ratios in Table 1. Many features difficult to see in wholemounted specimens. Body small, broadly oval, distinctly wider than long (Fig. 2). Tegument unarmed. Oral sucker transversely elongate oval, more or less terminal. Ventral sucker rounded to oval, larger than oral sucker, just-pre-equatorial. Prepharynx very short, dorsal to oral sucker. Pharynx large, oval. Oesophagus distinct, sinuous, thin-walled. Intestinal bifurcation dorsal to ventral sucker. Caeca blind, wide, divergent, almost or just reaching to testes. Testes two, irregular, symmetrical, separated in anterior hindbody. Cirrus sac claviform, reaches dorsally to ventral sucker. Internal seminal vesicle saccular. Pars prostatica short. Ejaculatory duct long, muscular. Genital atrium small. Genital pore median, about mid-way between suckers, but not clear in wholemounts. Ovary subglobular, just anterior to right testis, overlapping ventral sucker. Seminal receptacle uterine. Laurer���s canal opens dorsally in anterior hindbody. Mehlis��� gland not detected. Uterus reaching between testes, or just into post-testicular region, then passes over dorsal surface of ventral sucker. Eggs large, tanned, operculate, relatively few. Metraterm not distinct. Vitellarium follicular, in narrow arcuate lateral fields from pharynx or oesophagus to testicular level, mainly lateral to caeca, but encroaching dorsally. Excretory pore terminal. Vesicle short, I-shaped, reaches to uterus, not traced further. Remarks. This species appears to be a squat version of B. jolliveti, and differs in that it is always just wider than long, the tegument is wrinkled, the testes are lobate, and the caeca at their longest, only just reach to the testes (Table 1)., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on page 76, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628

Caudotestis ventichthysi n. sp. (Figs. 4–7) urn:lsid:zoobank.org:act: 96 F 5 B 7 AE-B 2 D 0-4 FBA- 9 A01- 13465 C 5 EF 4 D 8 Type host. Ventichthys biospeedoi Neilsen, Møller & Segonzac, 2006 (Ophidioformes: Ophidiidae), East-Pacific ventbrotula. Site. ? intestine (not given) Type locality. South East Pacific Rise, hydrothermal vent site Oasis, 17 ° 25.38 ’S, 113 ° 12.29 ’W; Biospeedo cruise, submersible Nautile, R/V Atalante, dive PL 1582, baited trap B 2, depth 2586 m, 22 April 2004. Biospeedo label: 1582 – 7. Type specimens. MNHM holotype 1 slide MNHN HEL 402, paratype 1 wholemount slide MNHN HEL 403, 1 section slide MNHN HEL 404; BMNH paratype 2013.12. 3.14. Etymology. This species is named after the host-genus. Description. Based on 4 specimens, one sectioned subsequent to study as wholemount. Measurements and ratios in Table 1. Body narrow fusiform, widest at level of ventral sucker (Figs. 4, 5). Tegument unarmed. Oral sucker cup-shaped, subterminal. Ventral sucker rounded, larger than oral sucker, equatorial. Prepharynx short. Pharynx oval, small. Oesophagus long, sinuous, thin-walled. Intestinal bifurcation distinctly anterior to ventral sucker. Caeca blind, divergent, narrow, reaching to posterior edge of ventral sucker. Testes oval to rounded, oblique to symmetrical, adjacent or contiguous, in mid-hindbody. Cirrus sac overlaps to mid- to posterior part of ventral sucker, seen clearly only in sections (Figure 6). Internal seminal vesicle saccular (Fig. 7). Pars prostatica thick-walled, with scattered gland-cells. Ejaculatory duct not clearly delimited from pars prostatica, short. Genital atrium small. Genital pore dextrally submedian, close to anterior edge of ventral sucker. Ovary oval to irregularly oval, anterior to dextral testis and separated by uterus. Seminal receptacle uterine. Laurer’s canal opens dorsally in anterior hindbody. Mehlis’ gland anterior to ovary. Uterus pre-testicular or overlaps to posterior margin of testes, runs dorsally along median line of ventral sucker. Eggs large, operculate, tanned, few. Distinct metraterm not detected. Vitellarium follicular, lateral fields reaching posterior pharynx or anterior oesophagus to anterior testis, interrupted at level of ventral sucker, approaching but not confluent in forebody. Excretory pore terminal. Vesicle shape I-shaped, reaches to ovary. Remarks. With its cirrus sac, uterine seminal receptacle, vitellarium not reaching posteriorly to the testes and non-lobate ovary, Caudotestis ventichthysi appears to be a fairly straightforward member of the genus Caudotestis and is distinguished as follows: Caudotestis nicolli differs in its longer caeca reaching to the testes or nearly so, longer cirrus sac with more elongate internal seminal vesicle, more extensive vitellarium not interrupted at the level of the ventral sucker, shorter forebody, larger eggs (80–120 x 48 –60 µm) and many other features. Caudotestis opisthorchis differs in its longer caeca overlapping the testes, longer cirrus sac with more elongate internal seminal vesicle, more extensive vitellarium not interrupted at the level of the ventral sucker, shorter forebody, larger eggs (74–108 x 37 –65 µm) and many other features. Caudotestis kerguelensis differs in its longer caeca reaching almost to the testes or overlapping them, more extensive vitellarium not interrupted at the level of the ventral sucker, larger eggs (82–92 x 45 –50 µm) and many other features. Caudotestis glacialis is similar to C. nicolli and differs from C. ventichthysi in its longer caeca reaching to the testes, longer cirrus sac, more extensive vitellarium not interrupted at the level of the ventral sucker, shorter forebody, larger eggs (93–115 x 51 –64 µm) and many other features. Caudotestis patagonensis is much larger and more elongate than C. ventichthysi. The forebody is short (about 24–25 % of body-length), the suckers are of similar size (width ratio 1: 0.89–1.38), there are big gaps between the ventral sucker and the ovary and the ovary and the anterior testis and the testes are in an elongate tail-like extension.

Subfamily Opecoelininae Gibson & Bray, 1984 urn:lsid:zoobank.org:act: 7947 AB04-DC04- 419 B- 8 E 3 F- 36 B 636 C 3874 B, Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on page 82, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628, {"references":["Gibson, D. I. & Bray, R. A. (1984) On Anomalotrema Zhukov, 1957, Pellamyzon Montgomery, 1957, and Opecoelina Manter, 1934 (Digenea: Opecoelidae), with a description of Anomalotrema koiae sp. nov. from North Atlantic waters. Journal of Natural History, 18, 949 - 964. http: // dx. doi. org / 10.1080 / 00222938400770831"]}

Caudotestis ventichthysi n. sp. (Figs. 4���7) urn:lsid:zoobank.org:act: 96 F 5 B 7 AE-B 2 D 0-4 FBA- 9 A01- 13465 C 5 EF 4 D 8 Type host. Ventichthys biospeedoi Neilsen, M��ller & Segonzac, 2006 (Ophidioformes: Ophidiidae), East-Pacific ventbrotula. Site. ? intestine (not given) Type locality. South East Pacific Rise, hydrothermal vent site Oasis, 17 �� 25.38 ���S, 113 �� 12.29 ���W; Biospeedo cruise, submersible Nautile, R/V Atalante, dive PL 1582, baited trap B 2, depth 2586 m, 22 April 2004. Biospeedo label: 1582 ��� 7. Type specimens. MNHM holotype 1 slide MNHN HEL 402, paratype 1 wholemount slide MNHN HEL 403, 1 section slide MNHN HEL 404; BMNH paratype 2013.12. 3.14. Etymology. This species is named after the host-genus. Description. Based on 4 specimens, one sectioned subsequent to study as wholemount. Measurements and ratios in Table 1. Body narrow fusiform, widest at level of ventral sucker (Figs. 4, 5). Tegument unarmed. Oral sucker cup-shaped, subterminal. Ventral sucker rounded, larger than oral sucker, equatorial. Prepharynx short. Pharynx oval, small. Oesophagus long, sinuous, thin-walled. Intestinal bifurcation distinctly anterior to ventral sucker. Caeca blind, divergent, narrow, reaching to posterior edge of ventral sucker. Testes oval to rounded, oblique to symmetrical, adjacent or contiguous, in mid-hindbody. Cirrus sac overlaps to mid- to posterior part of ventral sucker, seen clearly only in sections (Figure 6). Internal seminal vesicle saccular (Fig. 7). Pars prostatica thick-walled, with scattered gland-cells. Ejaculatory duct not clearly delimited from pars prostatica, short. Genital atrium small. Genital pore dextrally submedian, close to anterior edge of ventral sucker. Ovary oval to irregularly oval, anterior to dextral testis and separated by uterus. Seminal receptacle uterine. Laurer���s canal opens dorsally in anterior hindbody. Mehlis��� gland anterior to ovary. Uterus pre-testicular or overlaps to posterior margin of testes, runs dorsally along median line of ventral sucker. Eggs large, operculate, tanned, few. Distinct metraterm not detected. Vitellarium follicular, lateral fields reaching posterior pharynx or anterior oesophagus to anterior testis, interrupted at level of ventral sucker, approaching but not confluent in forebody. Excretory pore terminal. Vesicle shape I-shaped, reaches to ovary. Remarks. With its cirrus sac, uterine seminal receptacle, vitellarium not reaching posteriorly to the testes and non-lobate ovary, Caudotestis ventichthysi appears to be a fairly straightforward member of the genus Caudotestis and is distinguished as follows: Caudotestis nicolli differs in its longer caeca reaching to the testes or nearly so, longer cirrus sac with more elongate internal seminal vesicle, more extensive vitellarium not interrupted at the level of the ventral sucker, shorter forebody, larger eggs (80���120 x 48 ���60 ��m) and many other features. Caudotestis opisthorchis differs in its longer caeca overlapping the testes, longer cirrus sac with more elongate internal seminal vesicle, more extensive vitellarium not interrupted at the level of the ventral sucker, shorter forebody, larger eggs (74���108 x 37 ���65 ��m) and many other features. Caudotestis kerguelensis differs in its longer caeca reaching almost to the testes or overlapping them, more extensive vitellarium not interrupted at the level of the ventral sucker, larger eggs (82���92 x 45 ���50 ��m) and many other features. Caudotestis glacialis is similar to C. nicolli and differs from C. ventichthysi in its longer caeca reaching to the testes, longer cirrus sac, more extensive vitellarium not interrupted at the level of the ventral sucker, shorter forebody, larger eggs (93���115 x 51 ���64 ��m) and many other features. Caudotestis patagonensis is much larger and more elongate than C. ventichthysi. The forebody is short (about 24���25 % of body-length), the suckers are of similar size (width ratio 1: 0.89���1.38), there are big gaps between the ventral sucker and the ovary and the ovary and the anterior testis and the testes are in an elongate tail-like extension., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on pages 80-82, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628

Family Opecoelidae Ozaki, 1925 urn:lsid:zoobank.org:act: 47 AB 39 F 7 - 1 B 71 - 47 C 4 -A 918 -C 7601 ECF 5 AB 1, Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on page 74, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628, {"references":["Ozaki, Y. (1925) Preliminary notes on a trematode with anus. Journal of Parasitology, 12, 51 - 53."]}

Biospeedotrema biospeedoi n. sp. (Fig. 3) urn:lsid:zoobank.org:act: 8381 CFB 8 - 5 EA 9 - 4 FD 7 -A 9 DE-A 161 B 7 C0E 664 Type host. Thermichthys hollisi (Cohen, Rosenblatt & Moser) (Ophididiiformes: Bythitidae ), Hollis brosmekvabbe. Site. Small and large intestine. Type locality. South East Pacific Rise, hydrothermal vent site Hobbs, 17 �� 35.20 ���S, 113 �� 14.75 ���W; Biospeedo cruise, PL 1588, depth 2598 m, baited trap, 29 Apr. Biospeedo labels: 1588 ���3, 1588���6, 1588��� 8. Type specimens. MNHN holotype 1 slide MNHN HEL 391, paratypes 6 wholemount slides MNHN HEL 392 393 394 395 396 397, 4 section slides. MNHN HEL 398 399 400 401; BMNH paratypes 2013.12.3.7- 10. GenBank number. rDNA KF 733986. TABLE 1. Measurements and ratios of digeneans of hydrothermal vent fishes. Width as % of body-length 39.3���51.1 (43.6) 57.7���82.2 (67.9) 60.6 104���133 (121) 47.6���85.6 (71.6) 22.5���53.9 (37.2) ������ continued on the next page p TABLE 1. (Continued) Ventral sucker to ovary as % of body- 2.20���17.8 (7.41) 0���2.11 (0.40) 0 0���7.68 (2.21) 0���6.85 (1.88) 0���2.27 (0.75) length Etymology. This species is named after the Biospeedo expedition on which the material was collected Description. Based on 11 wholemounts and 2 sets of serial sections. Measurements and ratios in Table 1. Body small, oval to fusiform, widest at level of ventral sucker (Fig. 3). Tegument unarmed, but thrown into numerous denticulate annular folds. Oral sucker shallow cup-shaped, almost terminal. Ventral sucker rounded, larger than oral sucker, just pre-equatorial. Prepharynx short. Pharynx oval. Oesophagus long, thin-walled. Intestinal bifurcation dorsal to ventral sucker. Caeca blind, divergent, narrow, reaching just into hindbody or just to testis. Testes lobate, symmetrical, separated by uterus, in anterior hindbody. Cirrus sac elongate claviform, reaching into hindbody, proximal extremity in sinistral anterior hindbody, curves dorsally to ventral sucker. Internal seminal vesicle saccular. Pars prostatica narrow, surrounded by gland-cells. Ejaculatory duct? Genital atrium small. Genital pore median to slightly submedian, just anterior to ventral sucker. Ovary oval, between dextral testis and cirrus sac. Seminal receptacle uterine. Laurer���s canal opens dorsally in mid post-testicular region. Mehlis��� gland not clearly seen. Uterus mainly post-testicular but also intertesticular. Eggs large, operculate, tanned, few. Metraterm not clearly differentiated. Vitellarium follicular, follicles small, irregular, lateral fields reaching from anterior part of ventral sucker to anterior part of testes, confluent in narrow band dorsally to ventral sucker, lateral and dorsal to caeca. Excretory pore terminal, to dorsally subterminal. Vesicle I-shaped, reaches between testes to ovary. Remarks. In basic morphology this species is similar to B. jolliveti, but differs in being generally smaller. The vitelline fields are shorter, filling a smaller proportion of the body-length, and the cirrus sac is relatively larger (Table 1). The uterus tends to extend further posteriorly, relative to the testes. The morphological similarity between B. jolliveti and this species is also reflected in the molecular data; 1300 base pairs of lsrDNA differ only in two positions. Biospeedotrema parajolliveti differs in body-shape and the ratios of vitelline field length and ventral sucker size to body-length. The uterus tends to extend further posteriorly, relative to the testes., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on pages 76-80, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628

Biospeedotrema jolliveti n. sp. (Fig. 1) urn:lsid:zoobank.org:act: 5 F 4 B 8936 -D 162 - 4 F 77 - 9 D 80-240 E 32 DCAD 02 Type host. Ventichthys biospeedoi Neilsen, Møller & Segonzac (Ophidioformes: Ophidiidae), East-Pacific ventbrotula. Site. ? intestine (not given) Type locality. South East Pacific Rise, hydrothermal vent site Oasis, 17 ° 25.38 ’S, 113 ° 12.29 ’W; Biospeedo cruise, submersible Nautile, R/V Atalante, dive PL 1582, baited trap B 2, depth 2586 m, 22 April 2004, Biospeedo labels: 1582 –2, 1582– 5. Type specimens. MNHN holotype 1 slide MNHN HEL 373; paratypes 3 wholemount slides MNHN HEL 374 375 376, 5 section slides MNHN HEL 377 378 379 380 381; BMNH paratypes 2013.12.3.4- 6. GenBank numbers. rDNA KF 733985, rDNA KF 733988. Etymology. This species named after Dr. Didier Jollivet who led the Biospeedo expedition. Description. Based on 7 whole-mounts and 2 sets of serial sections specimens. Measurements and ratios in Table 1. Body small, broadly pyriform (Fig. 1). Tegument unarmed. Oral sucker subglobular, subterminal. Ventral sucker rounded to oval, much larger than oral sucker, centre just pre-equatorial. Prepharynx short, dorsal to oral sucker. Pharynx large, subglobular. Oesophagus distinct, thin-walled, long, sigmoid. Intestinal bifurcation about equatorial, dorsal to ventral sucker. Caeca blind, wide, divergent, reaching just past testes, not to posterior extremity. Testes two, oval to pyriform, transversely elongate, symmetrical to slightly oblique, in anterior hindbody. Cirrus sac claviform, reaches to about mid-ventral sucker. Internal seminal vesicle saccular. Pars prostatica long, narrow, surrounded by gland-cells. Ejaculatory duct short. Genital atrium distinct. Genital pore median to slightly dextrally submedian, immediately anterior to ventral sucker. Ovary oval to subtriangular, just anterior to or overlapping right testis, overlapping ventral sucker. Details of female proximal system disrupted in sections by eggs. Seminal receptacle uterine. Laurer’s canal opens dorsally at about level of posterior margin in ventral sucker. Mehlis’ gland ventral to ovary. Uterus reaching between testes, or just into post-testicular region, then passes sinistrally over dorsal surface of ventral sucker. Eggs large, tanned, operculate, relatively few. Metraterm not distinct. Vitellarium follicular, in lateral fields from pharynx or oral sucker to just into post-testicular region, narrow band confluent or approaching in region of testes, mainly lateral to caeca, but encroaching dorsally. Excretory pore terminal, in slight notch. Vesicle short, flattens into broad vesicle just posterior to testes. Remarks. The features differentiating this species from its congeners are discussed below.

Subfamily Stenakrinae Yamaguti, 1970 urn:lsid:zoobank.org:act:DB 62 B 9 C 8 - 5 C 6 A- 4096 - 95 CE-CFF 747 CB 4 B 69, Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on page 74, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628, {"references":["Yamaguti, S. (1970) Digenetic Trematodes of Hawaiian Fishes. Keigaku, Tokyo, 436 pp."]}

Buticulotrema thermichthysi n. sp. (Figs. 8���9) urn:lsid:zoobank.org:act:F 1 BC 29 FC- 883 D- 4 A 62-92 CC-C 54 B 6880 E039 Type host. Thermichthys hollisi (Cohen, Rosenblatt & Moser) (Ophididiiformes: Bythitidae ), Hollis brosmekvabbe. Site. Large intestine Type locality. South East Pacific Rise, hydrothermal vent site Hobbs, 17 �� 35.20 ���S, 113 �� 14.75 ���W; Biospeedo cruise, PL 1588, depth 2598 m, baited trap, 29 Apr. Biospeedo labels: 1588 ���3, 1588��� 7. Type specimens. MNHM holotype 1 slide MNHN HEL 405, paratypes 3 wholemount slides MNHN HEL 406 407, 6 section slides MNHN HEL 409 410 411 412 413 414; BMNH paratypes 2013.12.3.11- 13. GenBank numbers. rDNA KF 733984, rDNA KF 733987 Possible juvenile. Ventichthys biospeedoi, South East Pacific Rise, hydrothermal vent site Oasis, 17 �� 25.38 ���S, 113 �� 12.29 ���W; Biospeedo cruise, submersible Nautile, R/V Atalante, dive PL 1582, baited trap B 2, depth 2586 m, 22 April 2004. Biospeedo label: 1582 ��� 2 (juv.), MNHN HEL 408. Etymology. This species is named after the host-genus. Description. Based on 8 wholemounts and 4 sets of serial sections. Measurements and ratios in Table 1. Body elongate pyriform, widest in anterior hindbody (Fig. 8). Tegument unarmed. Oral sucker subglobular, subterminal. Ventral sucker rounded, larger than oral sucker, just pre-equatorial. Prepharynx dorsal to oral sucker. Pharynx subglobular, large. Oesophagus long, very thick-walled. Intestinal bifurcation dorsal to posterior part of ventral sucker. Caeca blind, narrow, reaching well into post-testicular region, but not to posterior extremity. Testes irregularly oval or pyriform, oblique, contiguous, in mid-hindbody. Cirrus sac absent. Seminal vesicle saccular, in anterior hindbody and overlapping ventral sucker, narrow distally to form coiled tube (Figs. 9). Pars prostatica long, with distinct field of gland-cells and fibrous tissue which may appear to be delimited in some views. Naked distal male duct (ejaculatory duct?) long, wall thickens slightly distally, no evidence of cirrus sac or everted cirrus. Genital atrium subglobular. Genital pore sinistral, beside mid- to posterior part of pharynx. Ovary oval to reniform, just anterior to dextral testis or more median. Female proximal system unclear, even in sections. Seminal receptacle saccular, presumably canalicular. Laurer���s canal not traced. Mehlis��� gland not clearly distinguished. Uterus pretesticular, intercaecal, passes to left of ventral sucker. Eggs large, operculate, tanned, numerous. Metraterm long, reaches from about mid-pars prostatica, fairly thin-walled. Vitellarium follicular, follicles large, field reaching from mid-pharynx to just into post-caecal region, not as far as posterior extremity, lateral and ventral to caeca, not confluent, although approaching in post-testicular zone. Excretory pore terminal, in slight indentation. Vesicle I-shaped, surrounded by prominent gland-cells posterior, reaches to testes. Remarks. The features distinguishing B. thermichthysi n. sp. from B. stenauchenus are the very long, very strongly muscular oesophagus, bifurcating dorsally to the posterior part of the ventral sucker, the long, narrow pars prostatica and distal male duct and the sinistral genital pore at the level of the pharynx. It is possible that the female proximal system has been misinterpreted, and the organ considered by us to be a canalicular seminal receptacle is a uterine seminal receptacle. If that were the case then this species would be close to the genus Pseudopecoelus Von Wicklen, 1946, a genus which has been frequently reported in deeper waters. None of the described species of Pseudopecoelus have a long, highly muscular oesophagus, reaching dorsally to the ventral sucker., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on pages 82-84, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628

Genus Buticulotrema Blend, Dronen & McEachran, 1993 urn:lsid:zoobank.org:act: 7331 E 813 - 20 EA- 4 F03-B 5 BC- 6215801 AE 23 D The only species in this genus Buticulotrema stenauchenus Blend, Dronen & McEachran, 1993, is reported from the macrourid fishes, the western softhead grenadier Malacocephalus occidentalis Goode & Bean, the common Atlantic grenadier Nezumia aequalis (G��nther) and the glasshead grenadier Hymenocephalus italicus Giglioli, in the deep-waters of the Gulf of Mexico (Blend et al. 1993, Blend in Klimpel et al. 2001)., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on page 82, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628, {"references":["Blend, C. K., Dronen, N. O. & McEachran, J. D. (1993) Buticulotrema stenauchenus n. gen., n. sp. (Digenea: Opecoelidae) from Malacocephalus occidentalis and Nezumia aequalis (Macrouridae) from the Gulf of Mexico. Journal of Parasitology, 79, 674 - 676. http: // dx. doi. org / 10.2307 / 3283601","Klimpel, S., Seehagen, A., Palm, H. W. & Rosenthal, H. (2001) Deep-water Metazoan Fish Parasites of the World. Logos, Berlin, 315 pp."]}

Subfamily Opecoelininae Gibson & Bray, 1984 urn:lsid:zoobank.org:act: 7947 AB04-DC04- 419 B- 8 E 3 F- 36 B 636 C 3874 B

Genus Biospeedotrema n. g. urn:lsid:zoobank.org:act: 6 A 819058 -BDC 2-4752 -A 4 DB- 9 A 47 A 5 EC 8 BDD Diagnosis. Body small, broadly pyriform to distinctly wider than long. Tegument unarmed, but may have annular wrinkles. Oral sucker subglobular to transversely oval, subterminal to terminal. Ventral sucker rounded to oval, larger than oral sucker, just pre-equatorial. Prepharynx short, dorsal to oral sucker. Pharynx large, subglobular to oval. Oesophagus distinct. Intestinal bifurcation about equatorial, dorsal to ventral sucker. Caeca blind, wide, divergent, reaching almost to or just past testes, not to posterior extremity. Testes two, oval, pyriform, may be lobate, symmetrical to slightly oblique, in anterior hindbody. Cirrus sac claviform, reaches dorsally to ventral sucker. Internal seminal vesicle saccular. Pars prostatica long, narrow, surrounded by gland-cells or indistinct. Ejaculatory duct short to long. Genital atrium distinct. Genital pore median to slightly dextrally submedian, immediately anterior to ventral sucker. Ovary subglobular, oval or subtriangular, just anterior to or overlapping right testis, overlapping ventral sucker. Seminal receptacle uterine. Laurer���s canal opens dorsally at about level of posterior margin in ventral sucker or anterior hindbody. Mehlis��� gland ventral to ovary. Uterus reaching between testes, or distinctly into post-testicular region. Eggs large, tanned, operculate, relatively few. Metraterm not distinct. Vitellarium follicular, in lateral fields from pharynx or oral sucker to just into post-testicular region, mainly lateral to caeca. Excretory pore terminal. Vesicle short, flattens into broad vesicle just posterior to testes. Type species Biospeedotrema jolliveti n. sp. Etymology. This new genus is named after the Biospeedo expedition on which the material was collected. Remarks. Biospeedotrema n. gen. is placed in the subfamily Stenakrinae as its members have a welldeveloped muscular cirrus sac and a uterine seminal receptacle. Cribb (2005) recognised six genera in the subfamily. The ovary is usually more or less globular to oval, but can be subtriangular. Utilising the key in Cribb (2005) the closest genus is Caudotestis Issaitschikov, 1928 (see below), but Biospeedotrema differs in the more or less symmetrical testicular configuration, with the uterus passing between the testes, sometimes distinctly into the post-testicular region. Of the four species of Stenakron Stafford, 1904 recognised by Cribb (2005), only one has symmetrical testes, namely Stenakron vitellosum (Manter, 1934) from the shortbeard codling Laemonema barbatulum Goode & Bean (Gadiformes, Moridae) in 538 +m depth (294 + fathoms) off Tortugas, Florida (Manter 1934). It has a trilobed ovary (a characteristic of the genus) and the uterus does not extend posteriorly to the testes. Biospeedotrema, Caudotestis and Stenakron are mainly parasites of deep-sea fishes (Klimpel et al. 2009)., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on pages 74-75, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628, {"references":["Cribb, T. H. (2005) Family Opecoelidae Ozaki, 1925. In: Jones, A., Bray, R. A. & Gibson, D. I. (Eds.), Keys to the Trematoda. Vol. 2. CABI Publishing and the Natural History Museum, Wallingford, pp. 443 - 531.","Manter, H. W. (1934) Some digenetic trematodes from deep-water fish of Tortugas, Florida. Papers from Tortugas Laboratory, 28, 257 - 345.","Klimpel, S., Busch, M. W., Kellermans, E., Kleinertz, S. & Palm, H. W. (2009) Metazoan Deep-Sea Fish Parasites. Heinrich Heine Universitat, Dusseldorf, 384 pp."]}

Genus Caudotestis Issaitschikow, 1928 urn:lsid:zoobank.org:act:ACF01C 2 B- 6130 - 4 D 82 -BC 8 E-CC 55 DF 11 B 2 B 7 The status of the genus Caudotestis was discussed by Cribb (2005), who restricted it to four species from cold and deep waters. A further species has been described since. These species are: The type-species Caudotestis nicolli Issaitschikow, 1928, is known from Cottidae and Anarhichadidae, Kara Sea and off the eastern coast of Canada (Issaitschikow 1928; Bray 1979). Caudotestis opisthorchis (Polyanski, 1955) is known from Agonidae, Cottidae, Gadidae, Hemitripteridae, Hexagrammidae, Liparidae, Psychrolutidae and Sebastidae, Barents Sea, East Kamchatka, Chukotsk, Bering Sea, Sea of Okhotsk, off British Columbia, off West Greenland and off the eastern coast of Canada (Shulman & Shulman-Albova 1953; Polyanski 1955; Strelkov 1960; Zhukov 1963; Mamaev 1965; Arai 1967, 1969; Brinkmann 1975; Bray 1979; Machida & Araki 1992; Shvetsova 1992, 1994). Caudotestis kerguelensis (Prudhoe & Bray, 1973) is known from Channichthyidae, Congiopodidae and Nototheniidae, off Heard Island, Kerguelen, and the Macquarie Islands (Prudhoe & Bray 1973; Zdzitowiecki & Pisano 1996). Caudotestis glacialis (Zdzitowiecki, 1989) (syn. Stenakron sp. of Prudhoe & Bray 1973) is known from Artedidraconidae, Bathydraconidae and Liparidae, Davis Sea, Bransfield Strait and the Weddell Sea (Prudhoe & Bray 1973; Zdzitowiecki 1989, 1993, 1997 a, 1997 b, 2002 a, 2000 b). Caudotestis patagonensis Cantatore, Lancia, Lanfranchi & Timi, 2012 is known from Congiopodidae, Patagonian Shelf, Argentina (Cantatore et al. 2012)., Published as part of Bray, R. A., Waeschenbach, A., Dyal, P., Littlewood, D. T. J. & Morand, S., 2014, New digeneans (Opecoelidae) from hydrothermal vent fishes in the south eastern Pacific Ocean, including one new genus and five new species, pp. 73-87 in Zootaxa 3768 (1) on page 80, DOI: 10.11646/zootaxa.3768.1.5, http://zenodo.org/record/285628, {"references":["Issaitschikow, I. M. (1928) [Contributions to the knowledge of parasitic helminths of some groups of vertebrates in the Russian Arctic]. Trudy Morskoi Nauchnyi Institut, Moskva, 3, 5 - 79 + pls I - II. [in Russian]","Cribb, T. H. (2005) Family Opecoelidae Ozaki, 1925. In: Jones, A., Bray, R. A. & Gibson, D. I. (Eds.), Keys to the Trematoda. Vol. 2. CABI Publishing and the Natural History Museum, Wallingford, pp. 443 - 531.","Bray, R. A. (1979) Digenea in marine fishes from the eastern seaboard of Canada. Journal of Natural History, 13, 399 - 431. http: // dx. doi. org / 10.1080 / 00222937900770331","Polyanski, Y. I. (1955) [Studies on the parasitology of the fish in the northern seas of the USSR. Parasites of fish of the Barents Sea. Trudy Zoologicheskogo Instituta, Leningrad, 19, 5 - 170. [in Russian; English translation 1966 Israel Program for Scientific Translations, 158 pp.]","Shulman, S. S. & Shulman-Albova, R. E. (1953) Parasites of Fish of the White Sea. Karelo-Finskii Institut Biologii, Akademii nauk SSSR, Leningrad, Moscow, 198 pp. [in Russian]","Strelkov, J. A. (1960) Endoparasitic worms of marine fishes of East Kamchatka. Trudy Zoologicheskogo Instituta, Leningrad, 28, 147 - 196. [in Russian, English summary]","Zhukov, E. V. (1963) [Parasite fauna of fishes in Chukotsk. Part II. Endoparasites of marine and freshwater fish]. Parazitologicheskii Sbornik, 21, 96 - 139. [in Russian]","Mamaev, Y. L. (1965) [Helminths of fish in the Bering Sea]. In: Leonov, V. A. et al., (Ed.) Parasitic Worms of Domestic and Wild Animals. Vladivostok: Akademii Nauk SSSR, pp. 168 - 188. [in Russian]","Arai, H. P. (1967) A preliminary report on a study of the parasites of marine fishes of Burke Channel, British Columbia. MS Report, Fisheries Research Board of Canada, 925, 1 - 26.","Arai, H. P. (1969) Preliminary report on the parasites of certain marine fishes of British Columbia. Journal of the Fisheries Research Board of Canada, 26, 2319 - 2337. http: // dx. doi. org / 10.1139 / f 69 - 226","Brinkmann, A. (1975) Trematodes from Greenland. Meddelelser om Gronland Udgivne af Kommissionen for Videnskabelige Undersogelser i Gronland, 205, 1 - 88.","Machida, M. & Araki, J. (1992) Some trematodes and cestodes from the Okhotsk Sea off Hokkaido, Japan. Memoirs of the National Science Museum, Tokyo, 25, 97 - 103.","Shvetsova, L. S. (1992) Pacific cod parasite infection rate. Possibility of utilization of this species. In: Gritsenko, O. F. et al., (Ed.) Living resources of the Pacific Ocean. VNIRO, Moscow, pp. 20 - 37. [in Russian]","Shvetsova, L. S. (1994) Rational Pacific cod utilization in connection with the parasitic invasion. Izvestiya of the Pacific Research Institute of Fisheries and Oceanography (TINRO). Vladivostok, 117, 115 - 131. [in Russian]","Prudhoe, S. & Bray, R. A. (1973) Digenetic trematodes from fishes. B. A. N. Z. Antarctic Research Expedition Reports, Series B (Zoology and Botany), 8, 199 - 225.","Zdzitowiecki, K. & Pisano, E. (1996) New records of Digenea infection elasmobranch and teleost fish off Heard Island (Kerguelen sub-region, sub-Antarctic). Archive of Fishery and Marine Research, 43, 265 - 272.","Zdzitowiecki, K. (1989) Morphologic and systematic data of 3 digenetic trematode species occurring in nototheniid fishes in the environs of the South Shetlands (Antarctic). Acta Parasitologica Polonica, 34, 319 - 324.","Zdzitowiecki, K. (1993) A re-examination of some Antarctic and subantarctic fish digeneans from the collection of the British Museum (Natural History). Acta Parasitologica, 38, 157 - 160.","Zdzitowiecki, K. (1997 a) Antarctic Digenea parasites of fishes. Synopsis of the Antarctic Benthos, 8, 1 - 156.","Zdzitowiecki, K. (1997 b) Digenea of fishes of the Weddell Sea. IV. Three opecoelid species of the genera Neolebouria, Helicometra and Stenakron. Acta Parasitologica, 42, 138 - 143.","Zdzitowiecki, K. (2002 a) Occurrence of Digenea in fishes of the family Artedidraconidae in the Weddell Sea and other areas of Antarctica. Acta Parasitologica, 47, 306 - 309.","Cantatore, D. M. P., Lancia, J. P., Lanfranchi, A. L. & Timi, J. T. (2012) A new species of Caudotestis (Digenea: Opecoelidae) parasitizing the horsefish, Congiopodus peruvianus (Scorpaeniformes: Congiopodidae) from the Patagonian Shelf, Argentina. Journal of Parasitology, 98, 1144 - 1147. http: // dx. doi. org / 10.1645 / ge- 3065.1"]}

Genus Buticulotrema Blend, Dronen & McEachran, 1993 urn:lsid:zoobank.org:act: 7331 E 813 - 20 EA- 4 F03-B 5 BC- 6215801 AE 23 D The only species in this genus Buticulotrema stenauchenus Blend, Dronen & McEachran, 1993, is reported from the macrourid fishes, the western softhead grenadier Malacocephalus occidentalis Goode & Bean, the common Atlantic grenadier Nezumia aequalis (Günther) and the glasshead grenadier Hymenocephalus italicus Giglioli, in the deep-waters of the Gulf of Mexico (Blend et al. 1993, Blend in Klimpel et al. 2001).

Redescription of Macrobothriotaenia ficta (Meggitt, 1931) Freze, 1965 (Figs. 1���22) Syns: Crepidobothrium fictum Meggitt, 1931; Proteocephalus fictus (Meggitt, 1931) Hughes, Baker & Dawson, 1941 Type and only host: Xenopeltis unicolor Reinwardt in Boie, 1827 (Ophidia: Xenopeltidae). Type locality: Rangoon (also Yangon), Burma (also Republic of the Union of Myanmar). Type material: BMNH 1932.3. 3.33. Site of infection: Intestine. Prevalence: 17 % (n = 6) in Hanoi, North Vietnam, 21.x. 2010; other data not available. Intensity of infection: one specimen found in X. unicolor from Hanoi, North Vietnam, 21. x. 2012; other data not available. Distribution: Indomalayan Region (South East Asia ��� Burma / Myanmar, Thailand, Vietnam). Re-description (based on 1 mature, newly collected specimen from Vietnam ��� MHNG-PLAT 75454, IPCAS C- 584 and 2 mature specimens from Thailand ��� BMNH 1976.4.13.14��� 15; measurements from the original description by Meggitt [1931] in square brackets): Proteocephalidae, Proteocephalinae. Testes, ovary, vitelline follicles and uterus medullary. Strobila acraspedote, with first immature proglottides much wider than long, last immature proglottides rectangular to slightly longer than wide, mature proglottides rectangular to longer than wide, pregravid and pregravid proglottides much longer than wide. Total length 51���62 mm (n = 3) [50 mm], maximum width 680 (n = 19) [800]. Proliferative zone (neck) narrow, 600���900 long and 165���420 wide (n = 3). Strobila consists of 132���220 proglottides: 102���183 immature (up to appearance of spermatozoa in vas deferens), 11���33 mature (up to appearance of eggs in uterus), 3���5 pregravid (up to appearance of hooks in oncospheres) and 13 gravid. Scolex highly modified, short and wide, 525���1290 long and 720���1685 wide (n = 4) [790���1000 wide, 590���830 dorso-ventrally], much wider than neck (n = 4), formed by 4 lobes (peduncles), each 400���775 wide, separated from each other by deep grooves (Figs. 1, 2, 11 ���13, 22). Each lobe contains unilocular, pincer-shaped sucker, 290���420 in diameter (n = 8), situated within cavity of each lobe, opening outside posterolaterally (Figs. 1 ���3, 11��� 13). Apical organ not observed, probably absent. Apical surface of scolex and upper non-adherent surface of suckers covered with long capilliform filitriches (Figs. 5, 6). Luminal surface of suckers covered with capilliform filitriches and few gladiate spinitriches (Fig. 7). Non-adherent surface of suckers covered with long capilliform filitriches and few gladiate spinitriches (Fig. 8). Posterior part of suckers and proliferation zone covered with long gladiate spinitriches (Figs. 9, 10). Longitudinal internal musculature well developed, formed by wide bundles of muscle fibres (Figs. 17���19). Ventral osmoregulatory canals thin-walled, wide, with secondary canals (Fig. 15); dorsal osmoregulatory canal thickwalled (Fig. 17���19); canals situated between testes and vitelline follicles, sometimes overlapping them (Fig. 20). Testes medullary, oval, 45���65 long and 30���55 wide (n = 19) [43���59 in diameter], relatively few, 43���63 in number (x = 51, n = 30, CV = 10 %) [51���61], in 1 layer, forming 3 well separated fields separated by uterine stem and terminal genitalia (Figs. 14���16): aporal field composed of 20���31 testes (n = 30) [28���31, but illustrated with as many as 35 on aporal side ��� Fig. 6 in Meggitt 1931], preporal field with 10���21 testes and postporal fields with 5���14 testes, both on poral side [23���30 on poral side]. Testes present also in last gravid proglottides (Fig. 16). Vas deferens strongly coiled, voluminous, reaching midline of proglottides, but not crossing it aporally (Figs 14, 15). Cirrus-sac elongate, thin-walled, long, 410���680 long [21���23 (?); see remarks below] and 75���130 [12���17? see remarks below] wide; length/width ratio 32���68 % (n = 17); cirrus-sac may reach midline of proglottides, length of cirrus-sac representing 34���48 % of width of proglottides (x = 40 %, n = 19) [33 %]. Cirrus muscular, unarmed, occupies 1 / 2 to 4 / 5 of cirrus-sac. Ejaculatory duct coiled (Fig. 20). Genital pore irregularly alternating, postequatorial, situated at 44���62 % (x = 52 %, n = 16, CV = 9 %) of proglottis length from anterior margin. Genital atrium narrow and deep (Fig. 14���16). Ovary bilobed, medullary, follicular, relatively short, length of ovary representing 10���19 % of length of proglottides (n = 30). Mehlis��� gland about 40���50 in diameter, representing 6���12 % of proglottis width. Relative ovarian size (percentage of ovarian size in relation to that of entire proglottis ��� see de Chambrier et al. 2012 for details) 4.1 % of proglottis size. Proximal (preovarian) part of vaginal canal directed anteriorly, slightly sinuous, turned almost perpendicularly to proglottis axis at level of genital pore (Figs. 14, 15); distal (terminal) part of vagina near genital pore with thickened wall, surrounded by large, elongate vaginal sphincter; 30���40 long and 25���30 wide (Fig. 20). Vagina posterior in 63 % (n = 43) to cirrus-sac, anterior in 37 %. Vitelline follicles medullary, few in number, small, 20���30 long and 10���20 wide, form 2 narrow bands on dorsal side of lateral margins (Figs. 14���19); band on poral sides separated by terminal genitalia to preporal and postporal group (Figs. 14 ���16, 20). Bands slightly widening towards posterior margin of proglottides, with more follicles posteriorly, never reaching their anterior and posterior margins (postporal follicles reach anterior margin of ovary or its middle part); length of bands represents only 63���91 % and 72���92 % of length of proglottides on poral and aporal side, respectively (n = 13; Figs. 14���16). Primordium of uterine stem medullary, already present in immature proglottides. Uterus medullary, with development of type 1 as defined by de Chambrier et al. (2004). Briefly, uterine stem present as undifferentiated longitudinal median concentration of chromophilic cells in immature proglottides. In mature proglottides, uterine stem straight, occupying almost entire length of proglottides (Fig. 14), reaching anterior margin, but not posterior margin, never crossing posteriorly ovarian isthmus. Lumen appears in first mature proglottides. Diverticula (lateral branches) formed when eggs appear in uterus. In pregravid proglottides, eggs mature quickly. Pregravid and gravid proglottides with 26���37 diverticula on each side (n = 12) [31���35]; diverticula occupy most space (63���70 % of proglottis width) in last gravid proglottides (Fig. 16). Eggs spherical, with three-layered embryophore and rounded digitiform projections on its external surface. Hyaline outer envelope (collapsed in whole mounts) 80���120 in diameter (n = 8); thick, round embryophore consisting of 3 layers; outer layer 33���39 (n = 8) in diameter, thicker than nuclei-containing envelope; oncospheres spherical, 12���14 wide (n = 8), with 6 hooklets 8���11 (n = 14) long (Fig. 21). Remarks: The original description of Macrobothriotaenia ficta (as Crepidobothrium fictum) was brief and contained only basic information on its morphology and very few measurements (Meggitt 1931). The scolex, the morphology of which distinguishes M. ficta from all other proteocephalidean tapeworms, was described just: ���Scolex... provided with four suckers sunk within four spherical lobes nearly constricted off from the true scolex���. Although Meggitt (1931) admitted in the description of C. fictum that ���This type of the scolex approximates more to the Phyllobothrium (....) type than to the Crepidobothrium ���, he placed his new species in the latter genus. The original description contained only 2 relatively simple figures of the scolex and a (probably mature) proglottis, the latter with an incorrect scale bar (0.05 mm instead of 0.5 mm). Gravid proglottides, terminal genitalia (cirrus-sac and the distal part of the vaginal canal), eggs and cross sections were not illustrated. However, both figures of Meggitt (1931) provided the information sufficient for characterization of the new species, which markedly differs from other proteocephalidean cestodes in its scolex morphology (Rego 1994). Based on the peculiar shape of the scolex, with suckers lying on four protruding parts of the scolex, Freze (1965) proposed a new genus, Macrobothriotaenia, to accommodate Meggitt���s (1931) species. In addition, M. ficta possesses other distinctive morphological characteristics, such as the dorsal position of the vitelline follicles, eggs with a three-layered embryophore covered with short, rounded projections, the postequatorial position of the genital pore, the extraordinarily long cirrus-sac, which can reach up to the midline of the proglottides, the fairly low number of testes (less than 65), etc. Based on the new information provided in this paper, the generic diagnosis of Macrobothriotaenia is amended: Macrobothriotaenia Freze, 1965: Proteocephalidae, Proteocephalinae. Worms of medium size (5���6 cm long); strobila acraspedote, proglottides variable in shape, from short and wide to rectangular to much longer than wide in mature, pregravid and gravid proglottides. Scolex highly modified, much wider than neck, formed by four lobes (peduncles), separated by deep grooves, bearing unilocular, pincer-shaped suckers opening outside posterolaterally. Internal longitudinal musculature well developed. Testes medullary, few in number, in 3 separate lateral fields (aporal, preporal and postporal), in 1 layer, present also in last gravid proglottides. Cirrus-sac elongate, large, may reach mid-line of proglottis. Vagina posterior to cirrus-sac, occasionally anterior. Vaginal sphincter present. Genital pores irregularly alternating, postequatorial. Ovary medullary, follicular, short. Vitelline follicles not numerous, forming two narrow lateral bands on dorsal side of proglottides, not reaching to their anterior and posterior margins; bands slightly widening posteriorly; poral band interrupted by terminal genitalia (cirrus-sac and vagina). Formation of uterus of type 1 (see de Chambrier et al. 2004). Eggs with three-layered embryophore and short, rounded projections on its external surface. Parasites of snakes of the family Xenopeltidae in South East Asia. Type and only species Macrobothriotaenia ficta (Meggitt, 1931) Freze, 1965. Molecular data and phylogeny New sequences for proteocephalidean taxa used in this study were deposited in GenBank under accession Nos. KC 785980 ���KC 786025 (see Table 1 for full details). Six gene sequences from four taxa were not characterized as a result of PCR failure or weak products unsuitable for sequencing; all other sequences (46 for 13 taxa) were aligned by gene and concatenated by taxon. The full alignment was comprised of 5423 positions (40 % lsrDNA, 32 % ssrDNA, 17 % rrnL, 11 % cox 1), of which 442 positions across the alignment could not be unambiguously aligned and were excluded prior to analysis. A multi-gene phylogenetic analysis places M. ficta in a clade of Ophiotaenia spp., where O. lapata Rambeloson, Ranaivoson & de Chambrier, 2012 from Madagascar forms the sister group to (M. ficta ��� Vietnam, O. bungari de Chambrier, Binh & Scholz, 2012 ��� Vietnam, Ophiotaenia sp. from Compsophis ��� Madagascar ((Ophiotaenia sp. from Antaresia maculosa��� Australia, Ophiotaenia sp. from Notechis scutatus��� Australia) (O. gallardi [Johnston, 1911] ��� Australia, O. longmani ��� Australia))) (Fig. 23). Within the ingroup, Crepidobothrium gerrardii (Baird, 1860) forms the earliest diverging taxon, followed by Thaumasioscolex didelphidis Ca��eda-Guzm��n, de Chambrier & Scholz, 2001 (Fig. 23), the latter forming the sister group to the above mentioned clade., Published as part of Scholz, Tom����, Chambrier, Alain De, Kuchta, Roman, Littlewood, D. Timothy J. & Waeschenbach, Andrea, 2013, Macrobothriotaenia ficta (Cestoda: Proteocephalidea), a parasite of sunbeam snake (Xenopeltis unicolor): example of convergent evolution, pp. 485-499 in Zootaxa 3640 (3) on pages 488-493, DOI: 10.11646/zootaxa.3640.3.12, http://zenodo.org/record/222893

Family Allocreadiidae Looss, 1902 urn:lsid:zoobank.org:act: 4 F 447 E01- 69 C 0-4 DDA- 9433 - 7 C 5383312979, Published as part of Bray, Rodney A., Foster, Garth N., Waeschenbach, Andrea & Littlewood, D Timothy J, 2012, The discovery of progenetic Allocreadium neotenicum Peters, 1957 (Digenea: Allocreadiidae) in water beetles (Coleoptera: Dytiscidae) in Great Britain, pp. 58-70 in Zootaxa 3577 on page 61, DOI: 10.5281/zenodo.213869, {"references":["Looss, A. (1902) Uber neue und bekannte Trematoden aus Seeschildroten. Nebst Erorterungen zur Systematik un Nomenclatur. Zoologische Jahrbucher. Abteilung fur Systematik Okologie und Geographie der Tiere, 16, 411 - 894."]}

Allocreadium neotenicum Peters, 1957 urn:lsid:zoobank.org:act: 549 F 3979 - 54 E 2-4 EFC-B 579-640042 B 4 F 24 B Synonyms: Pseudallocreadium neotenicum (Peters, 1957) Yamaguti, 1971; Allocreadium sp. of Peters (1955); allocreadiid trematode of Jackson (1958). Probable synonym: Allocreadium sp. of Crawford (1940). Hosts: Hydroporus rufifrons (M��ller) (Coleoptera: Dytiscidae); Agabus melanarius Aub�� (Coleoptera: Dytiscidae); Agabus paludosus (Fabricius) (Coleoptera: Dytiscidae). Site: Haemocoel. Localities: H. rufifrons, Staveley, Westmorland, 21 / 10 / 2010, Coll: G. N. Foster, SD 47649802 Ings Pond, Westmorland, 03/ 11 / 2011, SD 442984, Coll: G. N. Foster; Ings, Westmorland, SD44249838, 21/ 10 / 2010, Coll: G. N. Foster; a shallow marsh near Jordan Lane, Westmorland, SD635901, 24/03/ 2012, Coll: G. N. Foster; A. melanarius, Hindleap Warren, Ashdown Forest, East Sussex, Oct. 1968, Coll: R.B. Angus; A. paludosus, near St Andrews, Fife, October 1956. Coll.: D.J. Jackson. Vouchers: NHMUK 2012.9.28.1- 11. Description: Based mainly on 6 specimens from H. rufifrons, because specimens from A. melanarius are immature and the single specimen from A. paludosus is damaged. Measurements in Table 2. Body broadly oval, robust (Fig. 2). Tegument unarmed. Eye-spots discrete, lateral to oesophagus. Oral sucker rounded, subterminal, no stylet seen. Ventral sucker transversely oval, larger than oral sucker, in anterior third of body. Prepharynx not seen. Pharynx oval. Oesophagus short. Intestinal bifurcation in posterior forebody or dorsal to ventral sucker. Caeca wide, reaching just into post-testicular zone or to level of posterior testis. Testes 2, large oval, entire, but flattened on contiguous parts of margins, oblique in anterior hindbody. Cirrussac oval, in forebody and overlapping ventral sucker (Fig. 3). Internal seminal vesicle broad, long with globular and tubular parts, coiled. Pars prostatica not distinct, but a few large glandular cells surround junction of seminal vesicle and ejaculatory duct. Ejaculatory duct long, thick-walled. Genital atrium distinct, surrounded by large gland cells. Genital pore median, bifurcal, surrounded by wide sphincter. Ovary large, oval, entire, mostly dorsal to ventral sucker, may reach into hindbody. Seminal receptacle overlapping ovary and anterior testis, in dorsal plane. Laurer���s canal opens dorsally to left caecum at level of ovary (Fig. 4). Mehlis��� gland not clearly distinguished. Uterus pretesticular, ventral and sinistro-lateral to ovary, partly dorsal to ventral sucker. Vitellarium follicular; follicles large, irregular, tightly packed, anterior extent at level of ventral sucker, reaching to posterior extremity, confluent in post-testicular region, overlaps testes slightly, dorsal and ventral. Metraterm approaches genital atrium from left, about half length of cirrus-sac, with glandular sheath. Eggs large, tanned, operculate, numerous; not found in specimens from Agabus melanarius. Excretory pore terminal. Vesicle traced to anterior testis in immature specimens (Fig. 5), not seen in adults., Published as part of Bray, Rodney A., Foster, Garth N., Waeschenbach, Andrea & Littlewood, D Timothy J, 2012, The discovery of progenetic Allocreadium neotenicum Peters, 1957 (Digenea: Allocreadiidae) in water beetles (Coleoptera: Dytiscidae) in Great Britain, pp. 58-70 in Zootaxa 3577 on pages 61-66, DOI: 10.5281/zenodo.213869, {"references":["Peters, L. E. (1957) An analysis of the trematode genus Allocreadium Looss with the description of Allocreadium neotenicum sp. nov. from water beetles. Journal of Parasitology, 43, 136 - 142. doi: 10.2307 / 3274638.","Yamaguti, S. (1971) Synopsis of digenetic trematodes of vertebrates. Keigaku, Tokyo, Vol. I, 1074 pp.; Vol. II, 349 pp.","Peters, L. E. (1955) Morphology of the adult and the miracidium of a progenetic species of Allocreadium from water beetles of the family Dytiscidae. Journal of Parasitology, 41, Suppl., 36.","Jackson, D. J. (1958) Notes on some nematodes and trematodes infesting water beetles. Entomologist's Monthly Magazine, 94, 109 - 111.","Crawford, W. W. (1940) An unusual case of a sexually mature trematode from the body cavity of a diving beetle. Journal of Parasitology, 26, Suppl., 32."]}

Microplana kwiskea n. sp. Material examined: Holotype K9. Syntypes K1a to K8 (see above). All collected a few days prior to, and fixed on, 25 October 2004. Type locality: Bridge House, Kirby Wiske, Thirsk, Yorkshire, United Kingdom (Ordnance Survey Grid Reference: SE 375849). Etymology: The specific epithet “ kwiskea ” is an abbreviation of Kirby Wiske, the type locality. Description: Live animals (Fig. 1b, upper) are up to 2.5 cm long and about 2 mm in diameter at the widest point when extended. When contracted they are about 6 mm long and 3 mm wide (Fig. 1c, upper). They are cylindrical and taper towards the anterior and posterior ends both of which are blunt. They are a uniform mid-brown (khaki) colour, even ventrally. There is a creeping sole mid-ventrally in the form a narrow ridge. Observations on creeping animals through transparent tubes showed little muscular distortion of the sole suggesting that locomotion is primarily by means of the cilia of the sole. Two small eyes, visible as black spots, are present at the anterior end. Otherwise there are no markings or orifices visible on live specimens. Preserved specimens are up to 18 mm long, the anterior end particularly being contracted and curled ventrally (Fig. 16a & b). The mouth (pharyngeal aperture) is about 8 mm from the anterior end (44% of body length) and the gonopore is about 12 mm from the anterior end (67%), but these must be regarded as approximate due to contraction. The gonopore in the remaining preserved specimens (Fig. 16a & b) and in the sectioned specimens is raised and prominent with a slightly protruded penis, presumably from muscular contraction at fixation. The general anatomy is typically rhynchodemid. The two eyes of K8 have a diameter about 35 µm and length about 50 µm (Fig. 17b). The gut is triclad and there is a cylindrical pharynx. The pharynx of K2 is about 1.5 mm long and 0.5 mm in diameter (Fig. 17a). The wall of the pharynx is about 200 µm thick. The outer epithelium of the pharynx is about 2.5 µm thick and densely ciliated, the cilia being about 2.5 µm long. Inside this is a loose layer of radial and longitudinal muscle about 160 µm thick with an inner dense layer of circular and longitudinal muscle about 40 µm thick. The inner wall is not ciliated. The creeping sole (Fig. 17d & e) is distinct, narrow (0.5 mm wide, about one third of the total width) and raised as a mid-ventral ridge (0.25 mm high). The epidermis of the creeping sole is about 20 µm thick, has few, narrow, rhabdites and is densely ciliated with cilia about 5 µm long. Dorsal and lateral epithelium is about 30 µm thick with copious broad rhabdites but apparently devoid of cilia. Sub-epidermal muscle is indiscernable. Rhabdites are formed sub-epidermally, just inside the epidermis, about 50 µm from the outer surface. Inside the rhabdite-forming zone is a layer of parenchymal longitudinal muscle, about 50 µm thick dorsally and laterally, thickening ventrally over the creeping sole to about 100 µm. There are further longitudinal muscle bundles dorsal to the ventral nerve cords (Fig. 17d & e). The paired ventral nerve cords are joined by occasional transverse commissures (Fig. 17d & e). The epithelium of the intestinal diverticula is between 50 µm and 100 µm thick and contains numerous very small refractive granules. The paired ovaries of K8 are 500 µm from the anterior end (Fig. 17c). The paired ovo-vitelline ducts run posteriorly dorso-lateral to the ventral nerve cords. Testes lie ventrally between the intestinal diverticula. They are oval, about 100 µm by 70 µm. It is not possible to discern the sperm ducts in transverse sections of the anterior of the body, but posterior to the pharynx they run dorso-lateral to each ventral nerve cord in the vicinity of the ovo-vitelline ducts. The two sperm ducts broaden to about 30 µm in diameter just before they reach the penis to form sperm storage organs which are reflexed forwards (Fig. 18). Presumably these straighten on penis eversion. Each then narrows to about 10 µm external diameter, 5 µm internal diameter, enters the muscular base of the penis and the two ducts discharge separately but close together into the ejaculatory duct. The penis is long, about 1 mm, and the distal half is free in the antrum (Figs 16c & 18). For most of its length it is roughly parallel-sided, about 200 µm in diameter. The ejaculatory duct is broad, about 100 µm in diameter and has numerous internal short projections or folds. At its distal end (the last 250 µm) the penis narrows to a blunt point and the ejaculatory duct in the distal 100 µm is narrow, about 10 µm in diameter, and has a smooth internal wall. The free portion of the penis has a dense outer layer of circular muscle fibres about 15 µm thick. Transverse sections of the penis show it to consist mostly of dense circular muscle fibres with a thin layer of longitudinal muscle near the outer epithelium (Figs 16d & 19). The vagina is narrow and short, 320 µm long. The posterior end of the vagina receives the openings of the two ovo-vitelline ducts ventrally and the genito-intestinal duct dorsally. The genito-intestinal duct is clearly defined, strongly ciliated internally and has an internal diameter of 10 µm and external diameter of 30 µm for its entire length. It runs dorsally from the vagina, forwards and to the right to open into the right posterior intestinal diverticulum at about the level of the penis. The refractive granules of the digestive epithelium do not enter the genito-intestinal duct (Figs 19e & 20f). Specimen from Manchester. Collected 20 October 1985, Ecology Study Area, campus of the University of Manchester (Grid Reference: SJ846968). This specimen (Fig. 21) partially autodigested before it was fixed. It is not quite fully mature in that the penis is not very large, but the ejaculatory duct extends for most of its length and has internal folds or projections, similar to the above. The genito-intestinal duct is well-defined, narrow and runs directly to a digestive diverticulum on one side. Its identity was uncertain until compared with the above specimens from Kirby Wiske and we are confident that it is a specimen of M. kwiskea. Specimens from Shirdley Hill, Lancashire. Collected October 1992 at the site of population studies of Australoplana sanguinea (sensu Jones 1981) reported by Jones et al. (1998 and 2001) (Grid reference: SD360129). Two specimens have been sectioned (in addition to undoubted M. terrestris from the same site). Both have the same penis structure and direct genito-intestinal duct typical of M. kwiskea (Fig. 22) and again we are confident these are of that species. Discussion on Microplana kwiskea. The presence of a single pair of eyes and of a genito-intestinal connection means that these specimens are undoubtedly of the genus Microplana. There was little doubt about M. kwiskea being different from M. scharffi, the shape, size and colour being sufficient to distinguish the species. The different anatomy of the copulatory apparatus simply confirms this distinction. However, the size and shape of M. kwiskea is broadly similar to that of M. terrestris, though the colour is different, hence our original uncertainty. The molecular results are unequivocal, the specimens are of a different species to M. terrestris. The anatomy of the male ducts, penis and of the genito-intestinal duct are quite different in the two species. The expanded ejaculatory duct with internal projections (seminal vesicle of some authors) runs most of the length of the penis in M. kwiskea, but in M. terrestris it occupies only the basal half of the penis. In M. kwiskea, the genito-intestinal duct is of constant diameter for its entire length and it runs noticeably forwards to open into the intestine to one side (the right) of the penis. In M. terrestris the genito-intestinal duct broadens to a sinus-like cavity before narrowing to its opening or openings to the intestine which, though forwards of its origin, are somewhat dorsal and not as far forwards as in M. kwiskea. The pharyngeal opening (mouth) and gonopore are further apart in M. kwiskea than in M. terrestris. Apart from their different colour, probably the best discriminating feature between the two species when alive is the ventral creeping sole. This is best seen when the animals are viewed from the ventral side when placed in a transparent container. In M. terrestris the sole is wide, almost half the ventral width, hardly raised and almost white. In M. kwiskea the sole is narrow, ridged and virtually the same colour as the rest of the body., Published as part of Jones, Hugh D., Webster, Bonnie L., Littlewood, D. Timothy J. & Mcdonald, Jillian C., 2008, Molecular and morphological evidence for two new species of terrestrial planarians of the genus Microplana (Platyhelminthes; Turbellaria; Tricladida; Terricola) from Europe, pp. 1-38 in Zootaxa 1945 (1) on pages 23-32, DOI: 10.11646/zootaxa.1945.1.1, http://zenodo.org/record/5231281

Microplana kwiskea n. sp. Material examined: Holotype K9. Syntypes K1a to K8 (see above). All collected a few days prior to, and fixed on, 25 October 2004. Type locality: Bridge House, Kirby Wiske, Thirsk, Yorkshire, United Kingdom (Ordnance Survey Grid Reference: SE 375849). Etymology: The specific epithet “ kwiskea ” is an abbreviation of Kirby Wiske, the type locality. Description: Live animals (Fig. 1b, upper) are up to 2.5 cm long and about 2 mm in diameter at the widest point when extended. When contracted they are about 6 mm long and 3 mm wide (Fig. 1c, upper). They are cylindrical and taper towards the anterior and posterior ends both of which are blunt. They are a uniform mid-brown (khaki) colour, even ventrally. There is a creeping sole mid-ventrally in the form a narrow ridge. Observations on creeping animals through transparent tubes showed little muscular distortion of the sole suggesting that locomotion is primarily by means of the cilia of the sole. Two small eyes, visible as black spots, are present at the anterior end. Otherwise there are no markings or orifices visible on live specimens. Preserved specimens are up to 18 mm long, the anterior end particularly being contracted and curled ventrally (Fig. 16a & b). The mouth (pharyngeal aperture) is about 8 mm from the anterior end (44% of body length) and the gonopore is about 12 mm from the anterior end (67%), but these must be regarded as approximate due to contraction. The gonopore in the remaining preserved specimens (Fig. 16a & b) and in the sectioned specimens is raised and prominent with a slightly protruded penis, presumably from muscular contraction at fixation. The general anatomy is typically rhynchodemid. The two eyes of K8 have a diameter about 35 µm and length about 50 µm (Fig. 17b). The gut is triclad and there is a cylindrical pharynx. The pharynx of K2 is about 1.5 mm long and 0.5 mm in diameter (Fig. 17a). The wall of the pharynx is about 200 µm thick. The outer epithelium of the pharynx is about 2.5 µm thick and densely ciliated, the cilia being about 2.5 µm long. Inside this is a loose layer of radial and longitudinal muscle about 160 µm thick with an inner dense layer of circular and longitudinal muscle about 40 µm thick. The inner wall is not ciliated. The creeping sole (Fig. 17d & e) is distinct, narrow (0.5 mm wide, about one third of the total width) and raised as a mid-ventral ridge (0.25 mm high). The epidermis of the creeping sole is about 20 µm thick, has few, narrow, rhabdites and is densely ciliated with cilia about 5 µm long. Dorsal and lateral epithelium is about 30 µm thick with copious broad rhabdites but apparently devoid of cilia. Sub-epidermal muscle is indiscernable. Rhabdites are formed sub-epidermally, just inside the epidermis, about 50 µm from the outer surface. Inside the rhabdite-forming zone is a layer of parenchymal longitudinal muscle, about 50 µm thick dorsally and laterally, thickening ventrally over the creeping sole to about 100 µm. There are further longitudinal muscle bundles dorsal to the ventral nerve cords (Fig. 17d & e). The paired ventral nerve cords are joined by occasional transverse commissures (Fig. 17d & e). The epithelium of the intestinal diverticula is between 50 µm and 100 µm thick and contains numerous very small refractive granules. The paired ovaries of K8 are 500 µm from the anterior end (Fig. 17c). The paired ovo-vitelline ducts run posteriorly dorso-lateral to the ventral nerve cords. Testes lie ventrally between the intestinal diverticula. They are oval, about 100 µm by 70 µm. It is not possible to discern the sperm ducts in transverse sections of the anterior of the body, but posterior to the pharynx they run dorso-lateral to each ventral nerve cord in the vicinity of the ovo-vitelline ducts. The two sperm ducts broaden to about 30 µm in diameter just before they reach the penis to form sperm storage organs which are reflexed forwards (Fig. 18). Presumably these straighten on penis eversion. Each then narrows to about 10 µm external diameter, 5 µm internal diameter, enters the muscular base of the penis and the two ducts discharge separately but close together into the ejaculatory duct. The penis is long, about 1 mm, and the distal half is free in the antrum (Figs 16c & 18). For most of its length it is roughly parallel-sided, about 200 µm in diameter. The ejaculatory duct is broad, about 100 µm in diameter and has numerous internal short projections or folds. At its distal end (the last 250 µm) the penis narrows to a blunt point and the ejaculatory duct in the distal 100 µm is narrow, about 10 µm in diameter, and has a smooth internal wall. The free portion of the penis has a dense outer layer of circular muscle fibres about 15 µm thick. Transverse sections of the penis show it to consist mostly of dense circular muscle fibres with a thin layer of longitudinal muscle near the outer epithelium (Figs 16d & 19). The vagina is narrow and short, 320 µm long. The posterior end of the vagina receives the openings of the two ovo-vitelline ducts ventrally and the genito-intestinal duct dorsally. The genito-intestinal duct is clearly defined, strongly ciliated internally and has an internal diameter of 10 µm and external diameter of 30 µm for its entire length. It runs dorsally from the vagina, forwards and to the right to open into the right posterior intestinal diverticulum at about the level of the penis. The refractive granules of the digestive epithelium do not enter the genito-intestinal duct (Figs 19e & 20f). Specimen from Manchester. Collected 20 October 1985, Ecology Study Area, campus of the University of Manchester (Grid Reference: SJ846968). This specimen (Fig. 21) partially autodigested before it was fixed. It is not quite fully mature in that the penis is not very large, but the ejaculatory duct extends for most of its length and has internal folds or projections, similar to the above. The genito-intestinal duct is well-defined, narrow and runs directly to a digestive diverticulum on one side. Its identity was uncertain until compared with the above specimens from Kirby Wiske and we are confident that it is a specimen of M. kwiskea. Specimens from Shirdley Hill, Lancashire. Collected October 1992 at the site of population studies of Australoplana sanguinea (sensu Jones 1981) reported by Jones et al. (1998 and 2001) (Grid reference: SD360129). Two specimens have been sectioned (in addition to undoubted M. terrestris from the same site). Both have the same penis structure and direct genito-intestinal duct typical of M. kwiskea (Fig. 22) and again we are confident these are of that species. Discussion on Microplana kwiskea. The presence of a single pair of eyes and of a genito-intestinal connection means that these specimens are undoubtedly of the genus Microplana. There was little doubt about M. kwiskea being different from M. scharffi, the shape, size and colour being sufficient to distinguish the species. The different anatomy of the copulatory apparatus simply confirms this distinction. However, the size and shape of M. kwiskea is broadly similar to that of M. terrestris, though the colour is different, hence our original uncertainty. The molecular results are unequivocal, the specimens are of a different species to M. terrestris. The anatomy of the male ducts, penis and of the genito-intestinal duct are quite different in the two species. The expanded ejaculatory duct with internal projections (seminal vesicle of some authors) runs most of the length of the penis in M. kwiskea, but in M. terrestris it occupies only the basal half of the penis. In M. kwiskea, the genito-intestinal duct is of constant diameter for its entire length and it runs noticeably forwards to open into the intestine to one side (the right) of the penis. In M. terrestris the genito-intestinal duct broadens to a sinus-like cavity before narrowing to its opening or openings to the intestine which, though forwards of its origin, are somewhat dorsal and not as far forwards as in M. kwiskea. The pharyngeal opening (mouth) and gonopore are further apart in M. kwiskea than in M. terrestris. Apart from their different colour, probably the best discriminating feature between the two species when alive is the ventral creeping sole. This is best seen when the animals are viewed from the ventral side when placed in a transparent container. In M. terrestris the sole is wide, almost half the ventral width, hardly raised and almost white. In M. kwiskea the sole is narrow, ridged and virtually the same colour as the rest of the body.

Microplana groga n. sp. Material examined: Specimens labelled “ M. terrestris Font Groga ”, NHML 1998.2.9.3-5, deposited by Mateos et al. (1998). Three specimens, FG1 (3 slides), FG2 (3 slides), FG3 (4 slides). Type locality: “Font Groga”, Massis de Collserola (Sant Cugat del Vallès, Barcelona, Spain). Etymology: the specific epithet is after the type locality. Holotype: Specimen FG 1, three slides FG1.1, FG1.2 and FG 1.3, labelled Font Groga, M. Terrestris, NHML 1998.2.9.3. Co-types FG 2 (4 slides, NHML 1998.2.9.4) and FG 3 (4 slides, NHML 1998.2.9.5), similarly labelled. Mateos et al. (1998) identified five specimens as M. terrestris. Three were sectioned and are the specimens deposited above. A further two were sacrificed for nucleic acid extraction and one of these is the source of the GenBank sequence AF178318 (Carranza, pers. comm.) which is identical to that of our specimens of M. kwiskea (K2, K8, K9) (Fig. 2). The worms were described as “reddish to brown with black granules all over the surface: anterior end darker; creeping sole white” (Mateos et al. 1998). Representative sections of these three specimens are shown in Figs 23–26. The distance between the mouth (pharyngeal aperture) and the gonopore can be measured in all three specimens. In FG1 it is 1.4 mm, in FG2 it is 1.7 mm, in FG3 it is 1 mm. The pharynx is cylindrical, 1.1 mm long in FG2 and 0.9 mm in FG3 (it is reflexed in FG 1), and about 0.4 mm in diameter in all three specimens (Fig. 23a–d). The pharyngeal aperture is two-thirds along the pharyngeal cavity. The outer epithelium of the pharynx is densely ciliated, cilia about 2.5 µm long. The pharyngeal musculature consists of an outer thin, sub-epidermal layer of circular muscle about 10 µm thick, a loose layer of longitudinal and radial fibres about 130 µm thick and an inner, dense layer of mainly circular fibres, but with some longitudinal fibres, about 40 µm thick. The inner epithelium is not ciliated. Though there are no transverse sections of the whole body, it can be seen in longitudinal sections that there is no longitudinal muscle immediately sub-epidermal but that longitudinal muscle fibres are present 30 to 60 µm inside the epidemis (Fig. 23e). There are few longitudinal fibres dorsal to the ventral nerve cords. In all three specimens, the sperm ducts are exceptionally broadly expanded between the pharynx and the anterior end of the penis (Fig. 23e). The ducts narrow, internal diameter about 5 µm, enter the penis musculature where they are convoluted, and discharge separately into the base of the ejaculatory duct (which is labeled “seminal vesicle” by Mateos et al. 1998). The penis is muscular and free in the antrum for about three-quarters of its length. The penis musculature is quite dense with, from outer to inner: dense circular muscle about 15 µm thick: dense longitudinal muscle about 10 µm thick; more loose radial muscle about 100 µm thick; circular muscle about 10–20 µm thick around the ejaculatory duct. The ejaculatory duct is broad, about 70 µm in diameter, for most of the length of the penis but narrows at the distal end of the penis to about 10 µm. There are no internal projections or folds on the wall, though there are some possible secretory products. There is a short vagina which at its posterior end receives the two ovo-vitelline ducts ventrally and the genito-intestinal canal dorsally (Mateos et al. label this as “bursa copulatrix”). In FG1 and FG2 the single genito-intestinal canal opens into one of the digestive diverticula via a confused, plate-like opening, possibly with more than one opening (Figs 24, 25). In FG3 the genito-intestinal canal forks to become Y-shaped with each upper branch opening into a digestive diverticulum on the opposite side (Fig. 26). These openings are more distinct than in FG1 and FG2. Discussion of M. groga. The presence of a single pair of eyes and of a genito-intestinal connection means that these specimens also are of the genus Microplana. However, they are distinct from both M. terrestris and M. kwiskea. The penis of M. terrestris has a broad, spherical proximal ejaculatory duct with internal projections (“seminal vesicle” of many authors) and a narrow, distal, muscular penis papilla. The ejaculatory duct of M. groga lacks such a differentiated portion. The genito-intestinal canal of M. terrestris broadens to a sinuslike cavity before narrowing to its opening or openings to the intestine. That of FG1 and FG2 is narrow for its entire length. That of FG3 is narrow but forked. The penis of FG1, FG2 and FG3 are similar, so the significance of the forked genito-intestinal canal in FG3 is uncertain. Its openings into the intestine on either side are rather different from FG1 and FG2. The lack of a distinct layer of longitudinal muscle dorsal to the ventral nerve cord also distinguishes these specimens from M. terrestris and also from M. kwiskea. The cox1 results (Fig. 2) suggest these specimens are identical with M. kwiskea. However, Mateos et al. (1998) noted that their specimens had a white sole, but the sole of M. kwiskea is of the general body colour. The mouth and gonopore are much closer together in the Font Groga specimens (1–1.7 mm apart) than in M. kwiskea (4 mm apart). The internal surface of the ejaculatory duct appears smooth in the Font Groga specimens, unlike M. kwiskea in which there are internal folds or projections. The musculature of the penis of the Font Groga specimens (outer to inner circular, longitudinal, radial and circular fibres) is different from that of M. kwiskea (dense circular fibres with a longitudinal layer). In the Font Groga specimens the genito-intestinal canal is narrow for its entire length and in two of the three specimens the duct divides near its entry to the digestive diverticulum so that there are two or three openings into the diverticulum (Figs 24 & 25, and Fig. 12 of Mateos et al. 1998). FG3 has a forked, Y-shaped genito-intestinal canal (Fig. 26). These openings are posterior to the penis and antrum and somewhat dorsal. The genito-intestinal canal of M. kwiskea is narrow for its entire length and opens into the intestine on one side at the level of the penis via a distinct papillate opening. These differences lead us to conclude on morphological evidence that these specimens are of a different species to M. kwiskea. That is despite the apparently unambiguous identity of the cox1 sequence from the specimen AF178318. Thus they are a previously undescribed species, Microplana groga. How can the identical cox1 sequences reported above (Fig. 2, GenBank AF178318, K2, K8 and K9) be explained? There are two possibilities. Firstly, since different individuals were sectioned and sequenced by Mateos et al. (1998) there is the possibility that the sequenced individual (AF178318) was in fact not of the same species as FG1, FG2 and FG3. Secondly, although less likely, it is possible that the small fragment of cox1 characterized is identical between M. kwiskea and M. groga. Although widely popular as a ‘barcoding’ gene, partial cox1 is not always capable of discriminating between species with absolute accuracy (e.g. Meyer and Paulay 2005). Also, the fragment of cox1 characterized by Mateos et al. (1998) and herein, is approximately half the size of the fragment (~650 bp) suggested by Hajibabaei et al. (2007) for DNA barcoding. Whilst it remains possible that the fragment of cox1 used in the current study fails to differentiate between what are clearly distinct species (M. kwiskea and M. groga), there is a need to sequence new specimens of M. groga as detailed in this study; namely to section part of the worms for species verification and sequence the remaining tissues. If the sequences remain identical, or insufficiently different to separate them in a phylogeny, the possibility that the fragment of cox1 is incapable of differentiating these taxa remains and additional sequence data will be required for further verification.

Microplana terrestris (Müller 1774) Müller’s (1774) original description (Fig. 10) is minimal and none of his specimens remain. Müller did not specify any location or even country for the three specimens he examined, though he was Danish and worked in Denmark. Minelli (1977) designated a specimen from Denmark as neotype for the species but gave no figure or description of this specimen. Probably the best descriptions and diagrams of the copulatory apparatus of this species are by von Kennel (1882) and Bendl (1908; 1909), though there are differences apparent in the two Bendl diagrams. Microplana terrestris (Müller), neotype (see Minelli, 1977), NHMD, TUR-17.1-8 (8 slides LS anterior including pharynx); TUR-19.1-12 (12 slides LS posterior including copulatory apparatus). Labels read: “ Microplana terrestris (O.F. Müll.); det. Minelli 1976; Kildeskoven; I Ellestub; 6.X.1918; Hj. Ditlevsen ”. Google Earth shows two places with the name Kildeskoven, one near Fredericia (Jutland), the other near Kalundborg (Zealand). Figure 11 shows a series of sections through the copulatory apparatus of this specimen. Each sperm duct expands to form a sperm storage organ immediately before entering the muscular bulb of the penis. The sperm ducts discharge separately into the base of the penis. The penis consists of two distinct parts, a basal bulb which does not project into the antrum, and a tapering, muscular, penis papilla which projects into the antrum. The basal bulb has an external diameter about 0.4 mm and there are several broad internal projections into its cavity. This cavity is equivalent to the “seminal vesicle” of many authors, though it is uncertain to what extent it is glandular. The projecting penis papilla has a basal diameter of about 0.2 mm and is about 0.5 mm long. A narrow ejaculatory duct, diameter about 25 µm, runs from the bulb to the distal end of the penis through the muscular part. The vagina opens directly off the antrum, is narrow and about 0.3 mm long. The paired ovo-vitelline ducts open separately but close together into its posterior extremity ventrolaterally. Also opening at its posterior extremity, but from the dorsal side is the genito-intestinal connection. This opens into a broad, sinus-like, cavity which, in this specimen, has a single narrow connection antero-dorsally to an intestinal diverticulum. Rhynchodemus terrestris NHMW, 2831 M257. This collection consists of 151 slides of 20 specimens, some immature, from various European localities. The material originates from L von Graff and his collaborators at the University of Graz and some slides can be recognized as the source of illustrations in Graff (1899) and Bendl (1908, 1909). Some labels are apparently in Bendl’s handwriting. Figure 12a shows a section of a specimen from County Galway, Ireland. There is a spherical basal bulb with internal projections, and a short, conical penis papilla projecting into the antrum. The vagina is short and, as above, the two oviducts and broad genito-intestinal canal open into its posterior end. Transverse sections of another specimen (no locality on label) show the broad genito-intestinal canal and its opening into the intestinal diverticulum (Fig. 12b & c). Rhynchodemus carli Fuhrmann 1914. NHMG (transferred from the University of Neuchâtel). 5 slides. Minelli (1977) considered this to be a synonym of Microplana terrestris, but as Minelli comments, “the type specimen is of very little help in assessing the true nature of the species”. The specimen was 10.5 mm long (preserved) and uniform brown/yellow in colour. Sections of the copulatory apparatus are poor, incomplete and not clear enough for a definitive identification. The ovaries are only partially mature and no ovo-vitelline duct is visible in transverse sections. However, numerous testes are visible. Expanded sperm ducts with stored sperm are visible posteriorly, but the specimen was damaged and nothing of the penis or other parts of the copulatory apparatus is present. The colour of the specimen suggests that this may not have been a specimen of M. terrestris, but we cannot be certain of its identity. Microplana terrestris MNHNP, AO 38-49. Labelled “Madé C 1-3 ”. This specimen is one of those reported by Beachamp (1957) from Madeira. The penis is as described above (Fig. 13a) thus its identification is confirmed. The copulatory apparatus of two specimens of M. terrestris from Shirdley Hill, Lancashire (HDJ collection) are shown in Fig. 13, b & c. Microplana terrestris T1, T2, T4, T7 (see above). The copulatory apparatus of T1, T2 and T4 all have a basal penial bulb with internal projections, and a short muscular penis papilla (Figs. 14 & 15). The vagina opens directly from the antrum and the genito-intestinal canal is broad and sinus-like. There are two openings from the genito-intestinal canal into the intestine in T1 and T2 but only one in T4. Refractive granules in the digestive epithelium do not enter the genito-intestinal canal (Figs 14c, 15b, 15d). We are confident that our identification of these specimens as M. terrestris is accurate. Discussion of Microplana terrestris. The size, grey or black colour and broad white ventral creeping sole are quite good discriminating factors for live specimens of this species. However the colour may be non-uniform as in the granular specimens. The copulatory apparatus is distinctive, the penis having a spherical basal ejaculatory duct (seminal vesicle) with internal projections, and a distal, muscular penis papilla projecting into the antrum. The genito-intestinal duct opens via a broad, sinus-like chamber into a branch of the intestine via one or two openings. Both Graff (1899) and Bendl (1908; 1909) show that there may be more than one such connection. Minelli’s (1977) figure, which is after Bendl (1908), is generally accurate with the exception that the entry of the sperm ducts (labelled "deferens") into the penis is misleading. The figure in Ball & Reynoldson (1981) is generally accurate with the exception that the cavity in the basal bulb of the penis (labelled “seminal vesicle“) is shown with a smooth lining without any of the internal projections visible in all the above specimens., Published as part of Jones, Hugh D., Webster, Bonnie L., Littlewood, D. Timothy J. & Mcdonald, Jillian C., 2008, Molecular and morphological evidence for two new species of terrestrial planarians of the genus Microplana (Platyhelminthes; Turbellaria; Tricladida; Terricola) from Europe, pp. 1-38 in Zootaxa 1945 (1) on pages 16-19, DOI: 10.11646/zootaxa.1945.1.1, http://zenodo.org/record/5231281, {"references":["Muller, O. F. (1774) Vermium terrestrium et fluviatilium, seu animalium Infusorium, Helminthicorum, et Testaceorum, non marinorum, succincta historia. Vol I mi pars altera. Havniae et Lipsiae.","Minelli, A. (1977) A taxonomic review of the terrestrial planarians of Europe. Bolletino Zoologica, 44, 399 - 419.","Bendl, W. E. (1908) Beitrage zur Kenntnis des Genus Rhynchodemus. Zeitschrift fur wissenschaftliche Zoologie, 89, 525 - 554.","Bendl, W. E. (1909) Europaische Rhynchodemiden. I. Zeitschrift fur wissenschaftliche Zoologie, 92, 51 - 74.","Graff, L. von. (1899) Monographie der Turbellarien. II. Tricladida Terricola (Landplanarien). W. Engelmann, Leipzig.","Fuhrmann, O. (1914) Zwei neue Landplanarien aus der Schweiz. Revue Suisse de Zoologie, 22, 435 - 456.","Ball, I. R. & Reynoldson, T. B. (1981) British Planarians. Synopses of the British Fauna (New Series) No. 19, eds D. M. Kermack & R. S. K. Barnes. Published for the Linnean Society of London & The Estuarine and Brackish-Water Sciences Association by Cambridge University Press, Cambridge."]}

Some taxonomic groups are less amenable to mitochondrial DNA barcoding than others. Due to the paucity of molecular information of understudied groups and the huge molecular diversity within flatworms, primer design has been hampered. Indeed, all attempts to develop universal flatworm-specific COI markers have failed so far. We demonstrate how high molecular variability and contamination problems limit the possibilities for barcoding using standard COI-based protocols in flatworms. As a consequence, molecular identification methods often rely on other widely applicable markers. In the case of Monogenea, a very diverse group of platyhelminth parasites, and Rhabdocoela, representing one-fourth of all free-living flatworm taxa, this has led to a relatively high availability of nuclear ITS and 18S/28S rDNA sequences on GenBank. In a comparison of the effectiveness in species assignment we conclude that mitochondrial and nuclear ribosomal markers perform equally well. In case intraspecific information is needed, rDNA sequences can guide the selection of the appropriate (i.e. taxon-specific) COI primers if available. Walter A. Boeger (Universidade Federal do Parana, Brazil), Thierry Backeljau, Marc De Meyer and Kurt Jordaens (Joint Experimental Molecular Unit, Royal Belgian Institute of Natural Sciences/Royal Museum for Central Africa, Belgium), Filip A. M. Volckaert (University of Leuven, Belgium) and Niels Van Steenkiste (Hasselt University, Belgium/Fisheries and Oceans Canada) are gratefully acknowledged for their input into this research. We thank Gontran Sonet (Royal Belgian Institute of Natural Sciences, Belgium) for providing some of the R-scripts and the anonymous reviewers who commented on this manuscript. T.H. was, at the time of writing, a post-doctoral fellow of the Research Foundation - Flanders (FWO-Vlaanderen). M.P.M.V. was supported by KU Leuven - VES/12/005 and by Research Programme G.0553.10 of the Research Foundation - Flanders, and is currently funded by Czech Science Foundation project no. P505/12/G112 (ECIP - Centre of excellence). This research received support from the SYNTHESYS Project (http://www.synthesys.info/) which is financed by European Community Research Infrastructure Action under the FP7 Integrating Activities Programme. Diplectanid molecular analyses were supported by the "Service de Systematique Moleculaire" of the Museum national d'histoire naturelle (CNRS UMS 2700) and the network "Bibliotheque du Vivant" funded by the CNRS, the Museum national d'histoire naturelle, the INRA and the CEA (Genoscope).